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BODILY  CHANGES 
IN  PAIN,  HUNGER, 
FEAR    AND  RAGE 


/ 


BODILY  CHANGES 
IN  PAIN,  HUNGER, 
FEAR  AND  RAGE 


AX  ACCOUNT  OF  RECENT  RE- 
SEARCHES INTO  THE  FUNCTION 
OF     EMOTIONAL     EXCITEMENT 


BY 
WALTER  B.   CANNON 

GEORGE    HIGGI.VSON    PROFESSOR    OF    PHYSIOLOGY   IN 
HARVARD  UNIVERSITY 


NEW  YORK  AND  LONDON 
D.  AITLETON  AND   COMPANY 
1915 


Copyright,  1915,  by 
D.  APPLETON  AND   COMPANY 


y 


^1 


Printed  in  the  United  States  of  America 


TO  MY  COLLABORATORS  IN  THESE  RESEARCHES 

DANIEL    DE    LA    PAZ 
ALFRED    T.    SHOHL 
WADE   S.    WRIGHT 
ARTHUR    L.    WASHBURN 
HENRY    LYMAN 
LEONARD    B.    NICE 
CHARLES   M.    GRUBER 
HOWARD   OSGOOD 
HORACE    GRAY 
WALTER    L.    MENDENHALL 

WITH    PLEASANT    MEMORIES    OF   OUR 
WORK    TOGETHER 


PREFACE 

Fear,  rage  and  pain,  and  the  pangs  of  hunger  are  alt 
primitive  experiences  "which  human  beings  share  with 
the  lower  animals.  These  experiences  are  properly  classed 
as  among  the  most  powerful  that  determine  the  action 
of  men  and  beasts.  A  knowledge  of  the  conditions 
which  attend  these  experiences,  therefore,  is  of  general 
and  fundamental  importance  in  the  interpretation  of 
behavior. 

During  the  past  four  years  there  has  been  conducted, 
in  the  Harvard  Physiological  Laboratory,  a  series  of  in- 
vestigations concerned  with  the  bodily  changes  which 
occur  in  conjunction  with  pain,  hunger  and  the  major 
emotions.  A  group  of  remarkable  alterations  in  the 
bodily  economy  have  been  discovered,  all  of  which  can 
reasonably  be  regarded  as  responses  that  are  nicely 
adapted  to  the  individual's  welfare  and  preservation. 
Because  these  physiological  adaptations  are  interesting 
both  in  themselves  and  in  their  interpretation,  not  only 
to  physiologists  and  psychologists,  but  to  others  as  well, 
it  has  .seemed  worth  while  to  gather  together  in  con- 
venient form  the  original  accounts  of  the  experiments, 
which  have  been  published  in  various  American  medical 
and  physiological  journals.  I  have,  however,  attempted 
to  arrange  the  results  and  discussions  in  an  orderly 
and  consecutive  manner,  and  I  have  tried  also  to  elim- 

vii 


PREFACE 
viii 

inate  or  incidentally  to  explain  the  technical  terms,  so 
mate  or  mci  ^  understood  by   any 

that  the  exposition  will  be  easily  nnu  '     ^^_ 

intelligent  reader  even  though  not  trained  m  the  med 

"r  interest  in  the  conditions  —ng  pa^^^ 
h.3er  and  strong  emotional  states  was  stimulated  dur- 
•:;Z  clrse  of 'a  previous  series  of  -arche-nj^ 

Jtor  activities  of  the  al— ry  f^^det  th" 
of  these  researches  appeared  m  ^^ll^^f^^^  ^n 

reported  in  that  volume.  ^    ^    ^^^^^^ 


CONTENTS 


CHAPTER  I 

PAGES 

THE   EFFECT    OF   THE    EMOTIONS    ON    DIGESTION 

Emotions  favorable  to  normal  secretion  of  the  digestive 
juices — Emotions  unfavorable  to  normal  secretion  of 
the  digestive  juices — Emotions  favorable  and  un- 
favorable to  contractions  of  the  stomach  and  in- 
testines— The  disturbing  effect  of  pain  on  di- 
gestion      1-21 


CHAPTER  II 

THE  GENERAL  ORGANIZATION  OF  THE  VIS- 
CERAL  NERVES    CONCERNED    IN    EMOTIONS 

The  outlying  neurones — The  three  divisions  of  the  out- 
lying neurones — The  extensive  distribution  of  neu- 
rones of  the  "sympathetic"  or  thoracico-lurabar  di- 
vision and  their  arrangement  for  diffuse  action — The 
arrangement  of  neurones  of  the  cranial  and  sacral 
divisions  for  specific  action — The  cranial  division  a 
conserver  of  bodily  resources — The  sacral  division 
a  group  of  mechanisms  for  emptying — The  sympa- 
thetic division  antagonistic  to  both  the  cranial  and 
the  sacral — Neurones  of  the  sympathetic  division  and 
adrenal  secretion  have  the  same  action 22-39 


X  CONTENTS 

CHAPTER  III 

PAGES 

METHODS   OF   DEMONSTRATING   ADRENAL   SECRE- 
TION  AND  ITS   NERVOUS   CONTROL 

The  evidence  that  splanchnic  stimulation  induces  ad- 
renal secretion — The  question  of  adrenal  secretion 
in  emotional  excitement — The  method  of  securing 
blood  from  near  the  adrenal  veins — The  method  of 
testing  the  blood  for  adrenin 40-51 

CHAPTER  IV 

ADRENAL      SECRETION      IN      STRONG      EMOTIONS 
AND  PAIN 

The  evidence  that  adrenal  secretion  is  increased  in  emo- 
tional excitement — The  evidence  that  adrenal  secre- 
tion is  increased  by  "painful"  stimulation — Confirma- 
tion of  our  results  by  other  observers     ....     52-65 

CHAPTER  V 

THE    INCREASE    OF   BLOOD    SUGAR   IN    PAIN    AND 
GREAT   EMOTION 

Glycosuria  from  pain — Emotional  glycosuria — The  role 
of  the  adrenal  glands  in  emotional  glycosuria     .     66-80 

CHAPTER  VI 

IMPROVED   CONTRACTION    OF   FATIGUED    MUSCLE 
AFTER  SPLANCHNIC  STIMULATION  OF  THE 
ADRENAL   GLAND 

The  nerve-muscle  preparation — The  splanchnic  prepara- 
tion— The  effects  of  splanchnic  stimulation  on  the 
contraction  of  fatigued  muscle — The  first  rise  in  the 
muscle  record — The  prolonged  rise  in  the  muscle 
record — The  two  factors :  arterial  pressure  and  adre- 
nal  secretion 81-94 


N,l 


CONTENTS  xi 

CHAPTER   VII 

PAGES 

THE    EFFECTS    ON    CONTRACTION    OF    FATIGUED 
MUSCLE    OF    VARYING    THE    ARTERIAL 
BLOOD   PRESSURE 

The  effect  of  increasing  arterial  pressure — The  effect  of 
decreasing  arterial  pressure — An  explanation  of  the 
effects  of  varying  the  arterial  pressure — The  value 
of  increased  arterial  pressure  in  pain  and  strong 
emotion 95-109 

CHAPTER  VIII 

THE    SPECIFIC    ROLE    OF    ADRENIN    IN    COUNTER- 
ACTING THE   EFFECTS  OF  FATIGUE 

Variations  of  the  threshold  stimulus  as  a  measure  of 
irritability — The  method  of  determining  the  threshold 
stimulus — The  lessening  of  neuro-muscular  irrita- 
bility by  fatigue — The  slow  restoration  of  fatigued 
muscle  to  normal  irritability  by  rest — The  quick  res- 
toration of  fatigued  muscle  to  normal  irritability 
by  adrenin — The  evidence  that  the  restorative  ac- 
tion of  adrenin  is  specific — The  point  of  action  of 
adrenin  in  muscle 110-134 

CHAPTER  IX 

THE  HASTENING  OF  THE  COAGULATION  OF  BLOOD 
BY   ADRENIN 

The  graphic  method  of  measuring  the  coagulation  time 
— The  effects  of  subcutaneous  injections  of  adrenin — 
The  effects  of  intravenous  injections — The  hastening 
of  coagulation  by  adrenin  not  a  direct  effect  on  the 
blood       135-160 

CHAPTER  X 

THE  HASTENING  OF  COAGULATION  OF  BLOOD  IN 
PAIN   AND  GREAT   EMOTION 

Coagulation  hastened  by  splanchnic  stimulation — Co- 
agulation not  hastened  by  splanchnic  stimulation  if 


xii  CONTENTS 

PAGES 

the  adrenal  glands  are  absent — Coagulation  hast- 
ened by  "painful"  stimulation — Coagulation  hastened 
in  emotional  excitement 161-183 

CHAPTEK  XI 

THE  UTILITY  OF  THE  BODILY  CHANGES  IN  PAIN 
AND   GREAT    EMOTION 

The  reflex  nature  of  bodily  responses  in  pain  and  the 
major  emotions,  and  the  useful  character  of  re- 
flexes— The  utility  of  the  increased  blood  sugar  as 
a  source  of  muscular  energy — The  utility  of  in- 
creased adrenin  in  the  blood  as  an  antidote  to  the 
effects  of  fatigue — The  question  whether  adrenin 
normally  secreted  inhibits  the  use  of  sugar  in  the 
body — The  vascular  changes  produced  by  adrenin 
favorable  to  supreme  muscular  exertion — The  changes 
in  respiratory  function  also  favorable  to  great  effort 
— The  effects  produced  in  asphyxia  similar  to  those 
produced  in  pain  and  excitement — The  utility  of 
rapid  coagulation  in  preventing  loss  of  blood    .     184-214 

CHAPTEK  XII 

THE  ENERGIZING  INFLUENCE  OF  EMOTIONAL  EX- 
CITEMENT 

"Reservoirs  of  power" — The  excitements  and  energies  of 
competitive  sports — Frenzy  and  endurance  in  cere- 
monial and  other  dances — The  fierce  emotions  and 
struggles  of  battle — The  stimulating  influence  of 
witnesses  and  of  music — The  feeling  of  power    .   215-231 

CHAPTER  XIII 
THE  NATURE  OF  HUNGER 

Appetite  and  hunger — The  sensation  of  hunger — The 
theory  that  hunger  is  a  general  sensation — ^Weak- 
ness of  the  assumptions  underlying  the  theory  that 
hunger  is  a  general  sensation — Body  need  may  exist 
without  hunger — The  theory  that  hunger  is  of  gen- 


CONTENTS  xiii 

PAGES 

eral  origin  does  not  explain  the  quick  onset  and  the 
periodicity  of  the  sensation — The  theory  that  hunger 
is  of  general  origin  does  not  explain  the  local  refer- 
ence— Plunger  not  due  to  emptiness  of  the  stomach 
— Hunger  not  due  to  hydrochloric  acid  in  the  empty 
stomach — Hunger  not  due  to  turgescence  of  the  gas- 
tric mucous  membrane — Hunger  the  result  of  con- 
tractions— The  "empty"  stomach  and  intestines  con- 
tract— Observations  suggesting  that  contractions 
cause  hunger — The  concomitance  of  contractions  and 
hunger  in  man 232-266 

CHAPTER  XIV 

THE  INTERRELATIONS  OF  EMOTIONS 

Antagonism  between  emotions  expressed  in  the  sym- 
pathetic and  in  the  cranial  divisions  of  the  auto- 
nomic system — Antagonism  between  emotions  ex- 
pressed in  the  s^-mpathetic  and  in  the  sacral  di- 
visions of  the  autonomic  system — The  function  of 
hunger — The  similarity  of  visceral  effects  in  differ- 
ent strong  emotions  and  suggestions  as  to  its  psy- 
chological significance 267-284 

CHAPTER  XV 

ALTERNATIVE    SATISFACTIONS    FOR    THE    FIGHT- 
ING EMOTIONS 

Support  for  the  militarist  estimate  of  the  strength  of 
the  fighting  emotions  and  instincts — Growing  op- 
position to  the  fighting  emotions  and  instincts  as 
displayed  in  war — The  desirability  of  preserving  the 
martial  virtues — Moral  substitutes  for  warfare — Phy- 
sical substitutes  for  warfare — The  significance  of  in- 
ternational athletic  competitions 285-301 

A  LIST  OF  PUBLISHED  RESEARCHES  FROM  THE 
PHYSIOLOGICAL  LAP.ORATORY  IN  HARVARD 
rXIVEKSITY 302-303 

INDEX 305 


BODILY  CHAIN^GES   m  PAIN, 
HUI^GER,   FEAR  AND  RAGE 


CHAPTER    I 

THE    EFFECT    OF    THE    EMOTIONS    ON 
DIGESTION 

The  doctrine  of  human  development  from  sub- 
human antecedents  has  done  much  to  unravel  the 
complex  nature  of  man.  As  a  means  of  interpre- 
tation this  doctrine  has  been  directed  chiefly 
toward  the  solving  of  puzzles  in  the  peculiarities 
of  anatomical  structure.  Thus  arrangements  in 
the  human  body,  which  are  without  obvious  util- 
ity, receive  rational  explanation  as  being  vestiges 
of  parts  useful  in  or  characteristic  of  remote  an- 
cestors— jDarts  retained  in  man  because  of  age- 
long racial  inheritance.  This  mode  of  interpreta- 
tion has  proved  applicable  also  in  accounting  for 
functional  peculiarities.  Expressive  actions  and 
gestures — the  facial  appearance  in  anger,  for  ex- 
ample— observed  in  children  and  in  widely  dis- 
tinct races,  are  found  to  be  innate,  and  are  best 
explained  as  the  retention  in  human  beings  of 
responses  which  are  similar  in  character  in  lower 
animals. 

1 


2  BODILY    CHANGES 

From  this  point  of  view  biology  has  contributed 
much  to  clarify  our  ideas  regarding  the  motives 
of  human  behavior.  The  social  philosophies 
which  prevailed  during  the  past  century  either 
assumed  that  conduct  was  determined  by  a  cal- 
culated search  for  pleasure  and  avoidance  of  paui 
or  they  ascribed  it  to  a  vague  and  undefined 
faculty  named  the  conscience  or  the  moral  sense. 
Comparative  study  of  the  behavior  of  men  and 
of  lower  animals  under  various  circumstances, 
however,  especially  with  the  purpose  of  learning 
the  source  of  prevailing  impulses,  is  revealing  the 
inadequacy  of  the  theories  of  the  older  psychol- 
ogists. More  and  more  it  is  appearing  that  in 
men  of  all  races  and  in  most  of  the  higher  ani- 
mals, the  springs  of  action  are  to  be  found  in 
the  influence  of  certain  emotions  which  express 
themselves  in  characteristic  instinctive  acts. 

The  role  which  thes«  fundamental  responses  in 
the  higher  organisms  play  in  the  bodily  economy 
has  received  little  attention.  A^  a  realm  for  in- 
vestigation the  bodily  changes  in  emotional  ex- 
citement have  been  left  by  the  physiologists  to 
the  philosophers  and  psychologists  and  to  the 
students  of  natural  history.  These  students,  how- 
ever, have  usually  had  too  slight  experience  in 
the  detailed  examination  of  bodily  functions  to 
permit  them  to  follow  the  clues  which  superficial 
observation  might  present.     In  consequence  our 


EMOTIONS   AND    DIGESTION  3 

knowledge  of  emotional  states  lias  been  meagre. 
There  are,  of  course,  many  surface  manifesta- 
tions of  excitement.  The  contraction  of  blood 
vessels  with  resulting  pallor,  the  pouring  out  of 
"cold  sweat,"  the  stopping  of  saliva-flow  so  that 
the  "tongue  cleaves  to  the  roof  of  the  mouth,"  the 
dilation  of  the  pupils,  the  rising  of  the  hairs,  the 
rapid  beating  of  the  heart,  the  hurried  respira- 
tion, the  trembling  and  twitching  of  the  muscles, 
especially  those  about  the  lips — all  these  bodily 
changes  are  well  recognized  accompaniments  of 
pain  and  great  emotional  disturbance,  such  as 
fear,  horror  and  deep  disgust.  But  these  dis- 
turbances of  the  even  routine  of  life,  which  have 
been  commonly  noted,  are  mainly  superficial  and 
therefore  readily  observable.  Even  the  increased 
rapidity  of  the  heart  beat  is  noted  at  the  surface 
in  the  pulsing  of  the  arteries.  There  are,  how- 
ever, other  organs,  hidden  deep  in  the  body, 
which  do  not  reveal  so  obviously  as  the  struc- 
tures near  or  in  the  skin,  the  disturbances  of 
action  which  attend  states  of  intense  feeling. 
Special  methods  must  be  used  to  determine 
whether  these  deep-lying  organs  also  are  included 
in  the  complex  of  an  emotional*  agitation. 

*  In  the  use  of  the  term  "emotion"  the  meaning  here  is 
not  restricted  to  violent  affective  states,  but  includes  "feel- 
ings" and  other  affective  experiences.  At  times,  also,  in 
order  to  avoid  awkward  expressions,  the  term  is  used  in  the 
popular  manner,  as  if  the  "feeling"  caused  the  bodily  change. 


4  BODILY   CHANGES 

Among  tlie  organs  that  are  affected  to  an  im- 
portant degree  by  feelings  are  those  concerned 
with  digestion.  And  the  relations  of  feelings  to 
the  activities  of  the  alimentary  canal  are  of  par- 
ticular interest,  because  recent  investigations  have 
shown  that  not  only  are  the  first  stages  of  the 
digestive  process  normally  started  by  the  pleasur- 
able taste  and  smell  and  sight  of  food,  but  also 
that  pain  and  great  emotional  excitement  can 
seriously  interfere  with  the  starting  of  the  pro- 
cess or  its  continuation  after  it  has  been  started. 
Thus  there  may  be  a  conflict  of  feelings  and  of 
their  bodily  accompaniments — a  conflict  the  inter- 
esting bearing  of  which  we  shall  consider  later. 

Emotions  Favorable  to  Normal  Secretion  op  the 
Digestive  Juices 

The  feeliags  or  affective  states  favorable  to 
the  digestive  functions  have  been  studied  fruit- 
fully by  Pawlow,^  of  Petrograd,  through  in- 
genious experiments  on  dogs.  By  the  use  of  care- 
ful surgical  methods  he  was  able  to  make  a  side 
pouch  of  a  part  of  the  stomach,  the  cavity  of 
which  was  wholly  separate  from  the  main  cavity 
in  which  the  food  was  received.  This  pouch  was 
supplied  in  a  normal  manner  with  nerves  and 
blood  vessels,  and  as  it  opened  to  the  surface  of 
the  body,  the  amount  and  character  of  the  gastric 
juice    secreted   by    it   under   various    conditions 


EMOTIONS    AND    DIGESTION  5 

could  be  accurately  determined.  Secretion  by  that 
part  of  the  stomach  wall  which  was  included  in 
the  j^ouch  was  representative  of  the  secretory 
activities  of  the  entire  stomach.  The  arrange- 
ment was  particularly  advantageous  in  providing 
the  gastric  juice  unmixed  with  food.  In  some  of 
the  animals  thus  operated  upon  an  opening  was 
also  made  in  the  esophagus  so  that  when  the 
food  was  swallowed,  it  did  not  pass  to  the  stom- 
ach but  dropped  out  on  the  way.  All  the  pleas- 
ures of  eating  were  thus  experienced,  and  there 
was  no  necessity  of  stopping  because  of  a  sense 
of  fulness.  This  process  was  called  "sham  feed- 
ing." The  well-being  of  these  animals  was  care- 
fully attended  to,  they  lived  the  normal  life  of 
dogs,  and  in  the  course  of  months  and  years  be- 
came the  pets  of  the  laboratory. 

By  means  of  sham  feeding  Pawlow  showed  that 
the  chewing  and  swallowing  of  food  which  the 
dogs  relished  resulted,  after  a  delay  of  about  five 
minutes,  in  a  flow  of  natural  gastric  juice  from 
the  side  pouch  of  the  stomach — a  flow  which  per- 
sisted as  long  as  the  dog  chewed  and  swallowed 
the  food,  and  continued  for  some  time  after  eat- 
ing ceased.  Evidently  the  presence  of  food  in 
the  stomach  is  not  a  prime  condition  for  gastric 
secretion.  And  since  the  flow  occurred  only  when 
the  dogs  had  an  appetite,  and  the  material  pre- 
sented  to    them   was   agreeable,    the    conclusion 


V 


6  BODILY    CHANGES 

was    justified    that    this    was    a    true    psychic 
secretion. 

The  mere  sight  or  smell  of  a  favorite  food  may 
start  the  pouring  out  of  gastric  juice,  as  was 
noted  many  years  ago  by  Bidder  and  Schmidt  ^ 
in  a  hungry  dog  which  had  a  fistulous  opening 
through  the  body  wall  into  the  stomach.  This 
observation,  reported  in  1852,  was  confirmed  later 
by  Schiff  and  also  still  later  by  Pawlow.  That  ^ 
the  mouth  "waters"  with  a  flow  of  saliva  when 
palatable  food  is  seen  or  smelled  has  long  been 
such  common  knowledge  that  the  expression,  "It 
makes  my  mouth  water,"  is  at  once  recognized  as 
the  highest  testimony  to  the  attractiveness  of  an 
appetizing  dish.  That  the  stomach  also  "waters" 
in  preparation  for  digesting  the  food  which  is  to 
be  taken  is  clearly  proved  by  the  above  cited  ob- 
servations on  the  dog. 

The  importance  of  the  initial  psychic  secretion 
of  saliva  for  further  digestion  is  indicated  when, 
in  estimating  the  function  of  taste  for  the  pleas- 
ures of  appetite,  we  realize  that  materials  can 
be  tasted  only  when  dissolved  in  the  mouth  and 
thereby  brought  into  relation  with  the  taste  or- 
gans. The  saliva  which  "waters"  the  mouth  as- 
sures the  dissolving  of  dry  but  soluble  food  even 
when  it  is  taken  in  large  amount.  >. 

The  importance  of  the  initial  psychic  secretion 
of  gastric  juice  is  made  clear  by  the  fact  that  con- 


EMOTIONS   AND    DIGESTION  7 

tinuance  of  the  flow  of  this  juice  during  diges- 
tion is  provided  by  the  action  of  its  acid  or  its 
digestive  products  on  the  mucous  membrane  of 
the  pyloric  end  of  the  stomach,  and  that  secre- 
tion of  the  pancreatic  juice  and  bile  are  called 
forth  by  the  action  of  this  same  acid  on  the  mu- 
cous membrane  of  the  duodenum.  The  proper 
starting  of  the  digestive  process,  therefore,  is 
conditioned  by  the  satisfactions  of  the  palate,  and 
the  consequent  flow  of  the  first  digestive  fluids. 
The  facts  brought  out  experimentally  in  studies 
on  lower  animals  are  doubtless  true  also  of  man. 
Not  very  infrequently,  because  of  the  accidental 
swallowing  of  corrosive  substances,  the  esopha- 
gus is  so  injured  that,  when  it  heals,  the  sides 
grow  together  and  the  tube  is  closed.  Under 
these  circumstances  an  opening  has  to  be  made 
into  the  stomach  through  the  side  of  the  body  and 
then  the  individual  chews  his  food  in  the  usual 
manner,  but  ejects  it  from  his  mouth  into  a  tube 
which  is  passed  through  the  gastric  opening.  The 
food  thus  goes  from  mouth  to  stomach  through 
a  tube  outside  the  chest  instead  of  inside  the 
chest.  As  long  ago  as  1878,  Richet,^  who  had 
occasion  to  study  a  girl  whose  esophagus  was 
closed  and  who  was  fed  through  a  gastric  fistula, 
reported  that  whenever  the  girl  chewed  or  tasted 
a  highly  sapid  substance,  such  as  sugar  or  lemon 
juice,  while  the  stomach  was  empty,  there  flowed 


8  BODILY   CHANGES 

from  the  fistula  a  considerable  quantity  of  gastric 
juice.  A  number  of  later  observers  ^  have  had 
similar  cases  in  human  beings,  especially  in  chil- 
dren, and  have  reported  in  detail  results  which 
correspond  remarkably  with  those  obtained  in  the 
laboratory.  Hornborg'*  found  that  when  the 
little  boy  whom  he  studied  chewed  agreeable  food 
a  more  or  less  active  secretion  of  gastric  juice 
invariably  started,  whereas  the  chewing  of  an 
indifferent  substance,  as  gutta-percha,  was  fol- 
lowed by  no  secretion.  All  these  observations 
clearly  demonstrate  that  the  normal  flow  of  the 
first  digestive  fluids,  the  saliva  and  the  gastric 
juice,  is  favored  by  the  pleasurable  feelings 
which  accompany  the  taste  and  smell  of  food  dur- 
ing mastication,  or  which  are  roused  in  anticipa- 
tion of  eating  when  choice  morsels  are  seen  or 
smelled. 

These  facts  are  of  fundamental  importance  in 
the  serving  of  food,  especially  when,  through  ill- 
ness, the  appetite  is  fickle.  The  degree  of  dainti- 
ness with  which  nourishment  is  served,  the  little 
attentions  to  esthetic  details — the  arrangement 
of  the  dishes,  the  small  portions  of  food,  the 
flower  beside  the  plate — all  may  help  to  render 
food  pleasing  to  the  eye  and  savory  to  the  nos- 
trils and  may  be  the  deciding  factors  in  determin- 
ing whether  the  restoration  of  strength  is  to  be- 
gin or  not. 


EMOTIONS    AND    DIGESTION  9 

Emotions  Unfavorable  to  the  Normal  Secretion  of  the 
DiGESTRE  Juices 

The  conditions  favorable  to  proper  digestion 
are  wholly  abolished  when  un^Dleasant  feelings 
such  as  vexation  and  worry  and  anxiety,  or  great 
emotions  such  as  anger  and  fear,  are  allowed  to 
prevail.'  This  fact,  so  far  as  the  salivary  secre- 
tion is  concerned,  has  long  been  known.  The 
dry  mouth  of  the  anxious  person  called  upon  to 
speak  in  public  is  a  common  instance;  and  the 
"ordeal  of  rice,"  as  employed  in  India,  was  a  prac- 
tical utilization  of  the  knowledge  that  excitement 
is  capable  of  inhibiting  the  salivary  flow.  When 
several  persons  were  suspected  of  crime,  the  con- 
secrated rice  was  given  to  them  all  to  chew,  and 
after  a  short  time  it  was  spit  out  upon  the  leaf  of 
the  sacred  fig  tree.  If  anyone  ejected  it  dry,  that 
was  taken  as  proof  that  fear  of  being  discovered 
had  stopped  the  secretion,  and  consequently  he 
was  adjudged  guilty.^ 

What  has  long  been  recognized  as  true  of  the 
secretion  of  saliva  has  been  proved  true  also  of 
the  secretion  of  gastric  juice.  For  example, 
Hornborg  was  unable  to  confirm  in  his  little  pa- 
tient with  a  gastric  fistula  the  observation  by 
Pawlow  that  when  hunger  is  present  the  mere 
seeing  of  food  results  in  a  flow  of  gastric  juice. 
Hornborg  explained  the  difference  between  his 
and  Pawlow's  results  by  the  different  ways  in 


10  BODILY   CHANGES 

whicli  the  boy  and  tlie  dogs  faced  the  situation. 
When  food  was  shown,  but  withheld,  the  hungry 
dogs  were  all  eagerness  to  secure  it,  and  the  juice 
very  soon  began  to  flow.  The  boy,  on  the  con- 
trary, became  vexed  when  he  could  not  eat  at 
once,  and  began  to  cry;  then  no  secretion  ap- 
peared. Bogen  also  has  reported  the  instance  of 
a  child  with  closed  esophagus  and  gastric  fistula, 
who  sometimes  fell  into  such  a  passion  in  con- 
sequence of  vain  hoping  for  food  that  the  giving 
of  the  food,  after  the  child  was  calmed,  was  not 
followed  by  any  flow  of  the  secretion. 

The  inhibitory  influence  of  excitement  has  also 
been  seen  in  lower  animals  under  laboratory  con- 
ditions. Le  Conte*'  declares  that  in  studying 
gastric  secretion  it  is  necessary  to  avoid  all  cir- 
cumstances likely  to  provoke  emotional  reactions. 
In  the  fear  which  dogs  manifest  when  first 
brought  into  strange  surroundings  he  found  that 
activity  of  the  gastric  glands  may  be  completely 
suppressed.  The  suppression  occurred  even  if 
the  dog  had  eaten  freely  and  was  then  disturbed 
■ — as,  for  example,  by  being  tied  to  a  table.  When 
the  animals  became  accustomed  to  the  experi- 
mental procedure,  it  no  longer  had  an  inhibitory 
effect.  The  studies  of  Bickel  and  Sasaki'^  con- 
firm and  define  more  precisely  this  inhibitory 
effect  of  strong  emotion  on  gastric  secretion. 
They  observed  the  inhibition  on  a  dog  with  an 


EMOTIONS    AND    DIGESTION  II 

esophageal  fistula,  and  with  a  side  pouch  of  the 
stomach,  which,  as  in  Pawlow's  experiments, 
opened  only  to  the  exterior.  In  this  dog  Bickel 
and  Sasaki  noted,  as  Pawlow  had,  that  sham  feed- 
ing was  attended  by  a  copious  flow  of  gastric 
juice,  a  true  psychic  secretion,  resulting  from  the 
pleasurable  taste  of  the  food.  In  a  typical  in- 
stance the  sham  feeding  lasted  five  minutes,  and 
the  secretion  continued  for  twenty  minutes,  dur- 
ing which  time  66.7  cubic  centimeters  of  pure  gas- 
tric juice  were  produced. 

On  another  day  a  cat  was  brought  into  the 
presence  of  the  dog,  whereupon  the  dog  flew  into 
a  great  fury.  The  cat  was  soon  removed,  and 
the  dog  pacified.  Now  the  dog  was  again  given 
the  sham  feeding  for  five  minutes.  In  spite  of 
the  fact  that  the  animal  was  hungry  and  ate 
eagerly,  there  was  no  secretion  worthy  of  men- 
tion. During  a  period  of  twenty  minutes,  cor- 
responding to  the  previous  observation,  only  9 
cubic  centimeters  of  acid  fluid  were  produced,  and 
this  was  rich  in  mucus.  It  is  evident  that  in  the 
dog,  as  in  the  boy  observed  by  Bogen,  strong  emo- 
tions can  so  profoundly  disarrange  the  mechanisms 
of  secretion  that  the  pleasurable  excitation  which 
accompanies  the  taking  of  food  cannot  cause  the 
normal  flow. 

On  another  occasion  Bickel  and  Sasaki  started 
gastric  secretion  in  the  dog  by  sham  feeding,  and 


12  BODILY   CHANGES 

when  the  flow  of  gastric  juice  had  reached  a  cer- 
tain height,  the  dog  was  infuriated  for  five  min- 
utes by  the  presence  of  the  cat.  During  the  next 
fifteen  minutes  there  appeared  only  a  few  drops 
of  a  very  mucous  secretion.  Evidently  in  this 
instance  a  physiological  process,  started  as  an 
accompaniment  of  a  psychic  state  quietly  pleas- 
urable in  character,  was  almost  entirely  stopped 
after  another  psychic  state  violent  in  character. 
.  It  is  noteworthy  that  in  both  the  favorable  and 
unfavorable  results  of  the  emotional  excitement 
illustrated  in  Bickel  and  Sasaki's  dog  the  effects 
persisted  long  after  the  removal  of  the  exciting 
condition.  This  fact,  in  its  favorable  aspect, 
BickeP  was  able  to  confirm  in  a  girl  with 
esophageal  and  gastric  fistulas;  the  gastric  se- 
cretion long  outlasted  the  period  of  eating,  al- 
though no  food  entered  the  stomach.  The  in- 
fluences unfavorable  to  digestion,  however,  are 
stronger  than  those  which  promote  it.  And 
evidently,  if  the  digestive  process,  because  of 
emotional  disturbance,  is  for  some  time  inhibited, 
the  swallowing  of  food  which  must  lie  stagnant  in 
the  stomach  is  a  most  irrational  procedure.  If  a 
child  has  experienced  an  outburst  of  passion,  it 
is  well  not  to  urge  the  taking  of  nourishment  soon 
afterwards.  Macbeth's  advice  that  "good  diges- 
tion wait  on  appetite  and  health  on  both,"  is  now 
well-founded  physiology. 


EMOTIONS   AND    DIGESTION  13 

Other  digestive  glands  than  the  salivary  and 
the  gastric  may  be  checked  in  emotional  excite- 
ment. Eecently  Oechsler  ^  has  reported  that  in 
such  jDsychic  disturbances  as  were  shown  by 
Bickel  and  Sasaki  to  be  accompanied  by  sup- 
pressed secretion  of  the  gastric  juice,  the  secre- 
tion of  pancreatic  juice  may  be  stopped,  and  the 
flow  of  bile  definitely  checked.  All  the  means  of 
bringing  about  chemical  changes  in  the  food  may 
be  thus  temporarily  abolished. 

Emotions  Favorable  and  Unfavorable  to  the  Contractions 
OP  THE   Stomach  and  Intestines 

The  secretions  of  the  digestive  glands  and  the 
chemical  changes  wrought  by  them  are  of  little 
worth  unless  the  food  is  carried  onward  through 
the  alimentary  canal  into  fresh  regions  of  diges- 
tion and  is  thoroughly  exposed  to  the  intestinal, 
wall  for  absorption.  In  studying  these  mechani- 
cal aspects  of  digestion  I  was  led  to  infer  ^^  that 
just  as  there  is  a  psychic  secretion,  so  like- 
wise there  is  probably  a  "psychic  tone"  or  "psy- 
chic contraction"  of  the  gastro-intestinal  muscles 
as  a  result  of  taking  food.  For  if  the  vagus  nerve 
supply  to  the  stomach  is  cut  immediately  before 
an  animal  takes  food,  the  usual  contractions  of 
the  gastric  wall,  as  seen  by  the  Rontgen  rays,  do 
not  occur;  but  if  these  nerves  are  cut  after  food 
has  been  eaten  with  relish,  the  contractions  which 


14  BODILY   CHANGES 

have  started  continue  without  cessation.  The 
nerves  in  both  conditions  were  severed  under 
anesthesia,  so  that  no  element  of  pain  entered 
into  the  experiments.  In  the  absence  of  hunger, 
which  in  itself  provides  a  contracted  stomach/^ 
the  pleasurable  taking  of  food  may,  therefore,  be 
a  primary  condition  for  the  appearance  of  natural 
contractions  of  the  gastro-intestinal  canal. 

Again  just  as  the  secretory  activities  of  the 
stomach  are  unfavorably  influenced  by  strong 
emotions,  so  also  are  the  movements  of  the  stom- 
ach; and,  indeed,  the  movements  of  almost  the 
entire  alimentary  canal  are  wholly  stopped  dur- 
ing great  excitement.  In  my  earliest  observa- 
tions on  the  movements  of  the  stomach  ^^  I  had 
difficulty  because  in  some  animals  the  waves  of 
contraction  were  perfectly  evident,  while  in  others 
there  was  no  sign  of  activity.  Several  weeks 
passed  before  I  discovered  that  this  difference 
was  associated  with  a  difference  of  sex.  In  order 
to  be  observed  with  Rontgen  rays  the  animals 
were  restrained  in  a  holder.  Although  the  holder 
was  comfortable,  the  male  cats,  particularly  the 
young  males,  were  restive  and  excited  on  being 
fastened  to  it,  and  under  these  circumstances 
gastric  peristaltic  waves  were  absent ;  the  female 
cats,  especially  if  elderly,  usually  submitted  with 
calmness  to  the  restraint,  and  in  them  the  waves 
had  their  normal  occurrence.    Once  a  female  with 


EMOTIONS   AND   DIGESTION  15 

kittens  turned  from  lier  state  of  quiet  content- 
ment to  one  of  apjoarent  restless  anxiety.     The 
movements  of  the  stomach  immediately  stopped, 
the  gastric  wall  became  wholly  relaxed,  and  only 
after  the  animal  had  been  petted  and  began  to 
purr  did  the  moving  waves  start  again  on  their 
course.     By  covering  the  cat's  mouth  and  nose 
with  the  fingers  until  a  slight  distress  of  breath- 
ing is  produced,  the  stomach  contractions  can  be 
stopped  at  will.     In  the  cat,  therefore,  any  sign 
of  rage  or  fear,  such  as  was  seen  in  dogs  by  Le 
Conte  and  by  Bickel  and  Sasaki,  was  accompanied 
by   a   total   abolition   of   the   movements   of   the 
stomach.    Even  indications  of  slight  anxiety  may 
be  attended  by  complete  absence  of  the  churning 
waves.     In  a  vigorous  young  male  cat  I  have 
watched  the  stomach  for  more  than  an  hour  by 
means  of  the  Kontgen  rays,  and  during  that  time 
not  the  slightest  beginning  of  peristaltic  activity 
appeared;  yet  the  only  visible  indication  of  ex- 
citement  in  the   animal   was   a  continued   quick 
twitching  of  the  tail  to  and  fro.     What  is  true 
of  the  cat  I  have  found  true  also  of  the  rabbit, 
dog  and  guinea-pig  ^^ — very  mild  emotional  dis; 
turbances  are  attended  by  abolition  of  peristalsis. 
The  observations  on  the  rabbit  have  been  con- 
firmed by  Auer,^'*  who  found  that  the  handling 
of  the  animal  incidental  to  fastening  it  gently 
to    a   holder   stopped   gastric   peristalsis    for   a 


16  BODILY   CHANGES 

variable  length  of  time.  And  if  the  animal  was 
startled  for  any  reason,  or  struggled  excitedly, 
peristalsis  was  again  abolished.  The  observa- 
tions on  the  dog  also  have  been  confirmed;  Lom- 
mel  ^^  found  that  small  dogs  in  strange  sur- 
roundings might  have  no  contractions  of  the 
stomach  for  two  or  three  hours.  And  whenever 
the  animals  showed  any  indications  of  being  un- 
comfortable or  distressed,  the  contractions  were 
inhibited  and  the  discharge  of  contents  from  the 
stomach  checked. 

Like  the  peristaltic  waves  in  the  stomach,  the 
peristalsis  and  the  kneading  movements  (seg- 
mentation) in  the  small  intestine,  and  the  re- 
versed peristalsis  in  the  large  intestine  all  cease 
whenever  the  observed  animal  shows  signs  of 
emotional  excitement. 

There  is  no  doubt  that  just  as  the  secretory 
activity  of  the  stomach  is  affected  in  a  similar 
fashion  in  man  and  in  lower  animals,  so  likewise 
gastric  and  intestinal  peristaltic  waves  are 
stopped  in  man  as  they  are  stopped  in  lower  ani- 
mals, by  worry  and  anxiety  and  the  stronger 
affective  states.  The  conditions  of  mental  discord 
may  thus  give  rise  to  a  sense  of  gastric  inertia. 
For  example,  a  patient  described  by  Miiller^® 
testified  that  anxiety  was  always  accompanied  by 
a  feeling  of  weight,  as  if  the  food  remained  in 
the  stomach.    Every  addition  of  food  caused  an 


EMOTIONS   AND    DIGESTION  17 

increase  of  the  trouble.  Strong  emotional  states 
in  this  instance  led  almost  always  to  gastric  dis- 
tress, which  persisted,  according  to  the  grade  and 
the  duration  of  the  psychic  disturbance,  between 
a  half-hour  and  several  days.  The  patient  was 
not  hysterical  or  neurasthenic,  but  was  a  very 
sensitive  woman  deeply  affected  by  moods. 

The  feeling  of  heaviness  in  the  stomach,  men- 
tioned in  the  foregoing  case,  is  not  uncommonly 
complained  of  by  nervous  persons,  and  may  be 
due  to  stagnation  of  the  contents.  That  such 
stagnation  occurs  is  shown  by  the  following  in- 
stance. A  refined  and  sensitive  woman,  who  had 
had  digestive  difficulties,  came  with  her  husband 
to  Boston  to  be  examined.  They  went  to  a  hotel 
for  the  night.  The  next  morning  the  woman  ap- 
peared at  the  consultant's  office  an  hour  after 
having  eaten  a  test  meal.  An  examination  of  the 
gastric  contents  revealed  no  free  acid,  no  diges- 
tion of  the  test  breakfast,  and  the  presence  of  a 
considerable  amount  of  the  supper  of  the  pre- 
vious evening.  The  explanation  of  this  stagna- 
tion of  the  food  in  the  stomach  came  from  the 
family  doctor,  who  reported  that  the  husband 
had  made  the  visit  to  the  city  an  occasion  for  be- 
coming uncontrollably  drunk,  and  that  he  had 
by  his  escapades  given  his  wife  a  night  of  turbu- 
lent anxiety.  The  second  morning,  after  the 
woman   had  had   a  good   rest,   the  gastric  eon- 


18  BODILY   CHANGES 

tents  were  again  examined;  the  proper  acidity 
was  found,  and  the  test  breakfast  had  been  nor- 
mally digested  and  discharged. 

These  cases  are  merely  illustrative  and  doubt- 
less can  be  many  times  duplicated  in  the  experi- 
ence of  any  physician  concerned  largely  with  di- 
gestive disorders.  Indeed,  the  opinion  has  been 
expressed  that  a  great  majority  of  the  cases  of 
gastric  indigestion  that  come  for  treatment  are 
functional  in  character  and  of  jervous  origin.. 
It  is  the  emotional  element  that  seems  most  char- 
acteristic of  these  cases.  To  so  great  an  extent 
is  this  true  that  Rosenbach  has  suggested  that  as 
a  term  to  characterize  the  cause  of  the  distur- 
bances, "emotional"  dyspepsia  is  better  than 
"nervous"  dyspepsia.^'^ 

The  Disturbing  Effect  of  Pain  on  Digestion 

The  advocates  of  the  theory  of  organic  evolu- 
tion early  pointed  out  the  similarity  between  the 
bodily  disturbances  in  pain  and  in  the  major  emo- 
tions. The  alterations  of  function  of  internal  or- 
gans they  could  not  know  about.  The  general 
statement,  however,  that  pain  evokes  the  same 
changes  that  are  evoked  by  emotion,  is  true  also 
of  these  deep-lying  structures.  Wertheimer^^ 
proved  many  years  since  that  stimulation  of  a 
sensory  nerve  in  an  anesthetized  animal — such 
stimulation  as  in  a  conscious  animal  would  in- 


EMOTIONS   AND   DIGESTION  19 

duee  pain — qnicklj^  abolished  the  contractions  of 
the  stomach.  And  Netschaiev,  working  in  Paw- 
low's  ^^  laboratory,  showed  that  excitation  of 
the  sensory  fibres  in  the  sciatic  nerve  for  two  or 
three  minutes  resulted  in  an  inhibition  of  the 
secretion  of  gastric  juice  that  lasted  for  several 
hours.  Similar  effects  from  painful  experience 
have  been  not  uncommonly  noted  in  human  be- 
ings. Mantegazza,^^  in  his  account  of  the  physi- 
ology of  pain,  has  cited  a  number  of  such  ex- 
amples, and  from  them  he  has  concluded  that  pain 
interferes  with  digestion  by  lessening  appetite 
and  by  producing  various  forms  of  dyspepsia, 
with  arrest  of  gastric  digestion,  and  with  vomit- 
ing and  diarrhea.  The  expression,  "sickening 
pain"  is  testimony  to  the  power  of  strong  sensory 
stimulation  to  upset  the  digestive  processes  pro- 
foundly. Vomiting  is  as  likely  to  follow  violent 
pain  as  it  is  to  follow  strong  emotion.  A  "sick 
headache"  may  be,  indeed,  a  sequence  of  events 
in  which  the  pain  from  the  headache  is  primary, 
and  the  nausea  and  other  evidences  of  digestive 
disorder  are  secondary. 

As  the  foregoing  account  has  shown,  emotional 
conditions  or  "feelings"  may  be  accompanied  by 
quite  opposite  effects  in  the  alimentary  canal, 
some  highly  favorable  to  good  digestion,  some 
highly  disturbing.  It  is  an  interesting  fact  that 
the  feelings  having  those  antagonistic  actions  are 


20  BODILY   CHANGES 

typically  expressed  through  nerve  supplies  which 
are  correspondingly  opposed  in  their  influence 
on  the  digestive  organs.  The  antagonism  between 
these  nerve  supplies  is  of  fundamental  impor- 
tance in  understanding  not  only  the  operation  of 
conditions  favorable  or  unfavorable  to  digestion 
but  also  in  obtaining  insight  into  the  conflicts  of 
emotional  states.  Since  a  consideration  of  the 
arrangement  and  mode  of  action  of  these  nerves 
will  establish  a  firm  basis  for  later  analysis  and 
conclusions,  they  will  next  be  considered. 

EEFEEENCES 

^  Pawlow :  The  Work  of  the  Digestive  Glands,  London, 
1902. 

2  Bidder  and  Schmidt :  Die  Verdauungssaf  te  und  der 
Stoffwechsel,  Leipzig,  1852,  p.  35. 

2  Eichet :  Journal  de  I'Anatomie  et  de  la  Physiologic, 
1878,  xiv,  p.  170. 

*  See  Hornborg :  Skandinavisches  Archiv  f iir  Physiologie, 
1904,  XV,  p.  248.  Cade  and  Latarjet :  Journal  de  Physiologie 
et  Pathologie  Generale,  1905,  vii,  p.  221.  Bogen :  Archiv  fiir 
die  gesammte  Physiologie,  1907,  cxvii,  p.  156.  Lavenson: 
Archives  of  Internal  Medicine,  1909,  iv,  p.  271. 

^  Lea :    Superstition  and  Force,  Philadelphia,  1892,  p.  344. 

6Le  Conte:  La  Cellule,  1900,  xvii,  p.  291. 

''  Bickel  and  Sasaki :  Deutsche  medizinische  Wochen- 
schrift,  1905,  xxxi,  p.  1829. 

^  Bickel :  Berliner  klinische  Wochenschrif t,  1906,  xliii,  p. 
845. 

^Oechsler:  Internationelle  Beitrage  zur  Pathologie  und 
Therapie  der  Ernahrungstorungen,  1914,  v,  p.  1. 

^"  Cannon :  The  Mechanical  Factors  of  Digestion,  London 
and  New  York,  1911,  p.  200. 


EMOTIONS   AND    DIGESTION  21 

*^  Cannon  and  Washburn :  American  Journal  of  Physi- 
ology, 1912,  xxix,  p.  441. 

^-  Cannon :  The  American  Journal  of  Physiology,  1898,  i, 
p.  38. 

^^  Cannon :  American  Journal  of  Physiology,  1902,  vii, 
p.  xxii. 

^*  Auer :  American  Journal  of  Physiology,  1907,  xviii, 
p.  356. 

^^  Lommel :  Miinchener  medizinische  Wochenschrift, 
1903,  i,  p.  1634. 

^^  Miiller :  Deutsches  Archiv  f iir  klinische  Medicin,  1907, 
Ixxxix,  p.  434. 

^^Eosenbach:  Berliner  klinische  Wochenschrift,  1897, 
xxxiv,  p.  71 

18  ^Y'ertheimer:  Archives  de  Physiologic,  1892,  xxiv,  p. 
379. 

^®  Pawlow :  Loc.  cit.,  p.  56. 

^°  Mantegazza :  Fisiologia  del  Dolore,  Florence,  1880,  p. 
123. 


CHAPTER   II 

THE    GENERAL   ORGANIZATION    OF    THE 
VISCERAL  NERVES   CONCERNED   IN   EMOTIONS 

The  structures  of  the  alimentary  canal  which 
are  brought  into  activity  during  the  satisfactions 
of  appetite  or  are  checked  in  their  activity  during 
pain  and  emotional  excitement  are  either  the  se- 
creting digestive  glands  or  the  smooth  muscle 
which  surrounds  the  canal.  Both  the  gland  cells 
and  the  smooth-muscle  cells  differ  from  other 
tells  which  are  subject  to  nervous  influence — 
those  of  striated,  or  skeletal,  muscle — in  not  being 
directly  under  voluntary  control  and  in  being 
slower  in  their  response.  The  muscle  connected 
with  the  skeleton  responds  to  stimulation  within 
two  or  three  thousandths  of  a  second;  the  delay 
with  gland  cells  and  with  smooth  muscle  is  more 
likely  to  be  measured  in  seconds  than  in  fractions 
of  a  second. 

The  Outlying  Neurones 

The  skeletal  muscles  receive  their  nerve  supply 
direct  from  the  central  nervous  system,  i.  e.,  the 

22 


VISCERAL    NERVES  23 

nerve  fibres  distributed  to  these  muscles  are  parts 
of  neurones  whose  cell  bodies  lie  within  the  brain 
or  spinal  cord.  The  glands  and  smooth  muscles 
of  the  viscera,  on  the  contrary,  are,  so  far  as  is 
now  kno\Mi,  never  innervated  directly  from  the 
central  nervous  system.*  The  neurones  reaching 
out  from  the  brain  or  spinal  cord  never  come  into 
immediate  relation  with  the  gland  or  smooth- 
muscle  cells ;  there  are  always  interposed  between 
the  cerebrospinal  neurones  and  the  viscera  extra 
neurones  whose  bodies  and  processes  lie  wholly 
outside  the  central  nervous  system.  They  are 
represented  in  dotted  lines  in  Fig.  1.  I  have  sug- 
gested that  possibly  these  outlying  neurones  act 
as  "transformers,"  modifying  the  impulses  re- 
ceived from  the  central  source  (impulses  suited  to 
call  forth  the  quick  responses  of  skeletal  muscle), 
and  adapting  these  impulses  to  the  peculiar,  more 
slowly-acting  tissues,  the  secreting  cells  and  vis- 
ceral muscle,  to  which  they  are  distributed.^ 

The  outlying  neurones  typically  have  their  cell 
bodies  grouped  in  ganglia  (G's,  Fig.  1)  which,  in 
the  trunk  region,  lie  along  either  side  of  the  spinal 
cord  and  in  the  head  region  and  in  the  pelvic 
part  of  the  abdominal  cavity  are  disposed  near 
the  organs  which  the  neurones  supply.  In  some 
instances   these  neurones  lie  wholly  within  the 

*  The  special  case  of  the  adrenal  glands  will  be  considered 
later. 


Tear  gland 
Dilator  of  pupil 

Artery  of  salivary  gland 

Hair 

Surface  artery 
Sweat  gland 

Heart 

Hair 

Surface  artery 
^ — -^  Sweat  gland 

hy''^       Liver 

Stomach 
Visceral  artery 
Spleen 

Intestine 


Adrenal  gland 

Hair 

Surface  artery 

Sweat  gland 


Colon 

Bladder 

Rectum 

Artery  of  external 
genitals 

Figure  1. — Diagram  of  the  more  important  distributions  of  the 
autonomic  nervous  system.  The  brain  and  spinal  cord  are  repre- 
sented at  the  left.  The  nerves  to  skeletal  muscles  are  not  repre- 
sented. The  preganglionic  fibres  of  the  autonomic  system  are  in 
solid  hnes,  the  postganglionic  in  dash-lines.  The  nerves  of  the 
cranial  and  sacral  divisions  are  distinguished  from  those  of  the 
thoracico-lumbar  or  "sympathetic"  division  by  broader  Unes.  A  -\- 
mark  indicates  an  augmenting  effect  on  the  activity  of  the  organ; 
a  —  mark,  a  depressive  or  inhibitory  effect.  For  further  descrip- 
tion see  text. 


.VISCERAL   NERVES  25 

structure  which  they  innervate  ( see  e.  g.,  the  heart 
and  the  stomach,  Fig.  1).  In  other  instances  the 
fibres  passing  out  from  the  ganglia — the  so-called 
"postganglionic  fibres" — may  traverse  long  dis- 
tances before  reaching  their  destination.  The  in- 
nervation of  blood  vessels  in  the  foot  by  neurones 
whose  cell  bodies  are  in  the  lower  trunk  region 
is  an  example  of  this  extensive  distribution  of  the 
fibres. 

The  Three  Divisions  of  the  Outlying  Neurones 

As  suggested  above,  the  outlying  neurones  are 
connected  with  the  brain  and  spinal  cord  by 
neurones  whose  cell  bodies  lie  within  the  central 
nervous  organs.  These  connecting  neurones,  rep- 
resented in  continuous  lines  in  Fig.  1,  do  not  pass 
out  in  a  continuous'  series  all  along  the  cerebro- 
spinal axis.  Where  the  nerves  pass  out  from  the 
spinal  cord  to  the  fore  and  hind  limbs,  fibres  are 
not  given  off  to  the  ganglia.  Thus  these  connect- 
ing or  "preganglionic"  fibres  are  separated  into 
three  divisions.  In  front  of  the  nerve  roots  for 
the  fore  limbs  is  the  head  or  cranial  division,  be- 
tween the  nerve  roots  for  the  fore  limbs  and  those 
for  the  hind  limbs  is  the  trunk  division  (or  thorad- 
ico-lumbar  division,  or,  in  the  older  terminology, 
the  "sympathetic  system") ;  and  after  the  nerve 
roots  for  the  hind  limbs  the  sacral  division. 

This  system  of  outlying  neurones,  with  post- 


26  BODILY    CHANGES 

ganglionic  fibres  innervating  the  viscera,  and  with 
preganglionic  fibres  reaching  out  to  them  from 
the  cerebrospinal  system,  has  been  called  by 
Langley,  to  whom  we  are  indebted  for  most  of 
our  knowledge  of  its  organization,  the  autonomic 
nervous  system.^  This  term  indicates  that  the 
structures  which  the  system  supplies  are  not  sub- 
ject to  voluntary  control,  but  operate  to  a  large 
degree  independently.  As  we  have  seen,  a  highly 
potent  mode  of  influencing  these  structures  is 
through  conditions  of  pain  and  emotional  excite- 
ment. The  parts  of  the  autonomic  system — the 
cranial,  the  sympathetic,  and  the  sacral — ^have  a 
number  of  peculiarities  which  are  of  prime  im- 
portance in  accounting  for  the  bodily  manifesta- 
tions of  such  affective  states. 

The  ExTENsn^E  Distribution  of  Neurones  of  the  "Sympa- 
thetic" Division  and  Their  Arrangement  for  Diffuse 
Action 

The  fibres  of  the  sympathetic  division  differ 
from  those  of  the  other  two  divisions  in  being 
distributed  through  the  body  very  widely.  They 
go  to  the  eyes,  causing  dilation  of  the  pupils. 
They  go  to  the  heart  and,  when  stimulated,  they 
cause  it  to  beat  rapidly.  They  carry  impulses  to 
arteries  and  arterioles  of  the  skin,  the  abdominal 
viscera,  and  other  parts,  keeping  the  smooth  mus- 
cles of  the  vessel  walls  in  a  state  of  slight  con- 


VISCERAL   NERVES  27 

traction  or  tone,  and  thus  serving  to  maintain 
an  arterial  pressure  sufficiently  high  to  meet  sud- 
den demands  in  any  special  region;  or,  in  times 
of  special  discharge  of  impulses,  to  increase  the 
tone  and  thus  also  the  arterial  pressure.  They 
are  distributed  extensively  to  the  smooth  muscle 
attached  to  the  hairs;  and  when  they  cause  this 
muscle  to  contract,  the  hairs  are  erected.  They 
go  to  sweat  glands,  causing  the  outpouring  of 
sweat.  These  fibres  pass  also  to  the  entire  length 
of  the  gastro-intestinal  canal.  And  the  inhibi- 
tion of  digestive  activity  which,  as  we  have 
learned,  occurs  in  pain  and  emotional  states,  is 
due  to  impulses  which  are  conducted  outward  by 
the  splanchnic  nerves — the  preganglionic  fibres 
that  reach  to  the  great  ganglia  in  the  upper  abdo- 
men (see  Fig.  1) — and  thence  are  spread  by  post- 
ganglionic fibres  all  along  the  gut.^  They  in- 
nervate likewise  the  genito-urinary  tracts,  causing 
contraction  of  the  smooth  muscle  of  the  internal 
genital  organs,  and  usually  relaxation  of  the  blad- 
der. Finally  they  affect  the  liver,  releasing  the 
storage  of  material  there  in  a  manner  which  may 
be  of  great  service  to  the  bodj^  in  time  of  need. 
The  extensiveness  of  the  distribution  of  the  fibres 
of  the  sympathetic  division  is  one  of  its  most 
prominent  characteristics. 

Another  typical  feature  of  the  sympathetic  di- 
vision is  an  arrangement  of  neurones  for  diffuse 


28  BODILY   CHANGES 

discharge  of  the  nerve  impulses.    As  shown  dia- 
grammatically  in  Fig.  1,  the  preganglionic  fibres 
from   the   central   nervous    system   may   extend 
through  several  of  the  sympathetic  ganglia  and 
give  off  in  each  of  them  connections  to  cell  bodies 
of  the  outlying  neurones.    Although  the  neurones 
which  transmit  sensory  impulses  from  the  skin 
into  spinal  cord  have  similar  relations  to  nerve 
cells  lying  at  different  levels  of  the  cord,  the  op- 
eration in  the  two  cases  is  quite  different.     In 
the  spinal  cord  the  sensory  impulse  produces  di- 
rected and  closely  limited  effects,  as,  for  example, 
when  reflexes  are  being  evoked  in  a  "spinal"  ani- 
mal (i.  e.,  an  animal  with  the  spinal  cord  isolated 
from  the  rest  of  the  central  nervous  system),  the 
left  hind  limb  is  nicely  lifted,  in  response  to  a 
harmful  stimulus  applied  to  the  left  foot,  without 
widespread  marked  involvement   of  the   rest  of 
the  body  in  the  response.*     In  the  action  of  the 
sympathetic  division,  on  the  contrary,  the  con- 
nection of  single  preganglionic  fibres   with  nu- 
merous outlying  neurones  seems  to  be  not  at  all 
arranged  for  specific  effects  in  this  or  that  par- 
ticular region.    There  are,  to  be  sure,  in  different 
circumstances   variations   in   the   degree   of   ac- 
tivity of  different  parts;  for  example,  it  is  prob- 
able that  dilation  of  the  pupil  in  the  cat  occurs 
more  readily  than  erection  of  the  hairs.    It  may 
be  in  this  instance,  however,  that  specially  direct 


VISCERAL    NERVES  29 

pathways  to  tlie  eye  are  present  for  comnion  use 
in  non-emotional  states  (in  dim  light,  e.  g.),  and 
that  only  slight  general  disturbance  in  the  central 
nervous  system,  therefore,  would  be  necessary  to 
send  impulses  by  these  well-worn  courses.  Thus 
for  local  reasons  (dust,  e,  g.)  tears  might  flow 
from  excitation  of  the  tear  glands  by  sympathetic 
impulses,  although  other  parts  innervated  by  this 
same  division  might  be  but  little  disturbed.  "VVe 
have  no  means  of  voluntarily  wearing  these  path- 
ways, however,  and  both  from  anatomical  and 
physiological  evidence  the  neurone  relations  in 
the  sympathetic  division  of  the  autonomic  system 
seem  devised  for  widespread  diffusion  of  nervous 
impulses. 

The  Arrangement  op  Neurones  of  the  Cranial  and  Sacral 
Divisions  for  Specific  Action 

The  cranial  and  sacral  autonomic  divisions 
differ  from  the  sympathetic  in  having  only  re- 
stricted distribution  (see  Fig.  1).  The  third  cran- 
ial nerves  deliver  impulses  from  the  brain  to 
ganglia  in  which  lie  the  cell  bodies  of  neurones 
innervating  smooth  muscle  only  in  the  front  of 
the  eyes.  The  vagus  nerves  are  distributed  to 
the  lungs,  heart,  stomach,  and  small  intestine. 
As  shown  diagrammatically  in  Fig.  1,  the  out- 
lying neurones  in  the  last  three  of  these  organs 
lie  within  the  organs  themselves.  By  this  ar- 
rangement, although  the  preganglionic  fibres  of 


30  BODILY    CHANGES 

the  vagi  are  extended  in  various  directions  to 
structures  of  quite  diverse  functions,  singleness 
and  separateness  of  connection  of  the  peripheral 
organs  with  the  central  nervous  system  is  as- 
sured. The  same  specific  relation  between  effer- 
ent fibres  and  the  viscera  is  seen  in  the  sacral 
autonomic.  In  this  division  the  preganglionic 
fibres  pass  out  from  the  spinal  cord  to  ganglia 
lying  in  close  proximity  to  the  distal  colon,  the 
bladder,  and  the  external  genitals.  And  the  post- 
ganglionic fibres  deliver  the  nerve  impulses  only 
to  the  nearby  organs.  Besides  these  innervations 
the  cranial  and  sacral  divisions  supply  individual 
arteries  with  "dilator  nerves" — nerves  causing 
relaxation  of  the  particular  vessels.  Quite  typi- 
cally, therefore,  the  efferent  fibres  of  the  two 
terminal  divisions  of  the  autonomic  differ  from 
those  of  the  mid-division  in  having  few  of  the 
distributed  connections  characteristic  of  the  mid- 
division,  and  in  innervating  distinctively  the  or- 
gans to  which  they  are  distributed.  The  cranial 
and  sacral  preganglionic  fibres  resemble  thus  the 
nerves  to  skeletal  muscles,  and  their  arrangement 
provides  similar  possibilities  of  specific  and  sepa- 
rate action  in  any  part,  without  action  in  other 
parts. 

The  Cranial  Division  a  Conserver  op  Bodily  Kesources 

The    cranial    autonomic,    represented    by    the 

vagus  nerves,  is  the  part  of  the  visceral  nervous 


VISCERAL    NERVES  31 

system  concerned  in  the  psychic  secretion  of  the 
gastric  juice.  Pawlow  showed  that  when  these 
nerves  are  severed  psychic  secretion  is  abolished. 
The  cranial  nerves  to  the  salivary  glands  are  sim- 
ilarly the  agents  for  psj^chic  secretion  in  these 
organs,  and  are  known  to  cause  also  dilation  of  the 
arteries  supplying  the  glands,  so  that  during  ac- 
tivity the  glands  receive  a  more  abundant  flow  of 
blood.  As  previously  stated  (see  p.  13),  the  evi- 
dence for  a  psychic  tonus  of  the  gastro-intestinal 
musculature  rests  on  a  failure  of  the  normal  con- 
tractions if  the  vagi  are  severed  before  food  is 
taken,  in  contrast  to  the  continuance  of  the  con- 
tractions if  the  nerv^es  are  severed  just  after- 
wards. The  vagi  artificially  excited  are  well- 
known  as  stimulators  of  increased  tone  in  the 
smooth  muscle  of  the  alimentary  canal.  Aside 
from  these  positive  effects  on  the  muscles  of  the 
digestive  tract  and  its  accessory  glands,  cranial 
autonomic  fibres  cause  contraction  of  the  pupil 
of  the  eye,  and  slow^ing  of  the  heart  rate. 

A  glance  at  these  various  functions  of  the  cra- 
nial division  reveals  at  once  that  they  serve  for 
bodily  conservation.  By  narrowing  the  pupil  of 
the  eye  they  shield  the  retina  from  excessive 
light.  By  slowing  the  heart  rate,  they  give  the 
cardiac  muscle  longer  periods  for  rest  and  in- 
vigoration.  And  by  providing  for  the  flow  of 
saliva  and  gastric  juice  and  by  supplying  the  mus- 


32  BODILY    CHANGES 

cular  tone  necessary  for  contraction  of  the  ali- 
mentary canal,  they  prove  fundamentally  essen- 
tial to  the  processes  of  proper  digestion  and 
absorption  by  which  energy-yielding  material  is 
taken  into  the  body  and  stored.  To  the  cranial 
division  of  the  visceral  nerves,  therefore,  belongs 
the  qniet  service  of  building  up  reserves  and  forti- 
fying the  body  against  times  of  need  or  stress. 

The  Sacral  Division  a  Group  of  Mechanisms  for  Emptying 

Sacral  autonomic  fibres  cause  contraction  of  the 
rectum  and  distal  colon  and  also  contraction  of 
the  bladder.  In  both  instances  the  effects  result 
reflexly  from  stretching  of  the  tonically  con- 
tracted viscera  by  their  accumulating  contents. 
No  affective  states  precede  this  normal  action  of 
the  sacral  division  and  even  those  which  accom- 
pany or  follow  are  only  mildly  positive ;  a  feeling 
of  relief  rather  than  of  elation  usually  attends 
the  completion  of  the  act  of  defecation  or  mic- 
turition— though  there  is  testimony  to  the  con- 
trary. 

The  sacral  autonomic  fibres  also  include,  how- 
ever, the  nervi  erigentes  which  bring  about  en- 
gorgement of  erectile  tissue  in  the  external  geni- 
tals. According  to  Langley  and  Anderson^  the 
sacral  nerves  have  no  effect  on  the  internal  gen- 
erative organs.  The  vasa  deferentia  and  the 
seminal    vesicles    whose    rhythmic    contractions 


VISCERAL    NERVES  33 

mark  the  acme  of  sexual  excitement  in  the  male, 
and  the  uterus  whose  contractions  in  the  female 
are  probably  analogous,  are  supplied  only  by 
lumbar  branches — part  of  the  sympathetic  divi- 
sion. These  branches  also  act  in  opposition  to 
the  nervi  erigentes  and  cause  constriction  of  the 
blood  vessels  of  the  external  genitals.  The  sexual 
orgasm  involves  a  high  degree  of  emotional  ex- 
citement; but  it  can  be  rightly  considered  as  es- 
sentially a  reflex  mechanism ;  and,  again  in  this 
instance,  distention  of  tubules,  vesicles,  and  blood 
vessels  can  be  found  at  the  beginning  of  the  in- 
cident, and  relief  from  this  distension  at  the  end. 
Although  distention  is  the  commonest  occasion 
for  bringing  the  sacral  division  into  activity  it  is 
not  the  only  occasion.  Great  emotion,  such  as  is 
accompanied  by  nervous  discharges  via  the  sym- 
pathetic division,  may  also  be  accompanied  by  dis- 
charges via  the  sacral  fibres.  The  involuntary 
voiding  of  the  bladder  and  lower  gut  at  times  of 
violent  mental  stress  is  well-kno^\^l.  Veterans  of 
wars  testify  that  just  before  the  beginning  of  a 
battle  many  of  the  men  have  to  retire  temporarily 
from  the  firing  line.  And  the  power  of  sights 
and  smells  and  libidinous  thoughts  to  disturb  the 
regions  controlled  by  the  nervi  erigentes  proves 
that  this  part  of  the  autonomic  system  also  has 
its  peculiar  affective  states.  The  fact  that  one 
part  of  the   sacral   division,   e.  g.,   the   distribu- 


34  BODILY   CHANGES 

tion  to  the  bladder,  may  be  in  abeyance,  while 
another  part,  e.  g.,  the  distribution  to  the  rectum, 
is  active,  illustrates  again  the  directive  discharge 
of  impulses  which  has  been  previously  described 
as  characteristic  of  the  cranial  and  sacral  portions 
of  the  autonomic  system. 

Like  the  cranial  division,  the  sacral  is  engaged 
in  internal  service  to  the  body,  in  the  performance 
of  acts  leading  immediately  to  greater  comfort. 

The  Sympathetic  Division  Antagonistic  to  Both  the 
Cranial  and  the  Sacral 

As  indicated  in  the  foregoing  description  many 
of  the  viscera  are  innervated  both  by  the  cranial 
or  sacral  part  of  the  autonomic  and  by  the  sym- 
pathetic. When  the  mid-part  meets  either  end- 
part  in  any  viscus  their  effects  are  antagonistic. 
Thus  the  cranial  supply  to  the  eye  contracts  the 
pupil,  the  sympathetic  dilates  it;  the  cranial 
slows  the  heart,  the  sympathetic  accelerates  it; 
the  sacral  contracts  the  lower  part  of  the  large 
intestine,  the  sympathetic  relaxes  it;  the  sacral 
relaxes  the  exit  from  the  bladder,  the  sym- 
pathetic contracts  it.  These  opposed  effects 
are  indicated  in  Fig.  1  by  +  for  contraction,  ac- 
celeration or  increased  tone;  and  by  -  for  inhibi- 
tion, relaxation,  or  decreased  tone.* 

*  The  vagus  nerve,  when  artificially  stimulated,  has  a  pri- 
mary, brief  inhibitory  effect  on  the  stomach  and  small  intes- 
tine; its  main  function,  however,  as  already  stated,  is  to  pro- 


VISCERAL    NER\^S  35 

Sherrington  has  demonstrated  that  the  setting 
of  skeletal  muscles  in  opposed  groups  about  a 
joint  or  system  of  joints — as  in  flexors  and  ex- 
tensors— is  associated  with  an  internal  organiza- 
tion of  the  central  nervous  system  that  provides 
for  relaxation  of  one  group  of  the  opposed  mus- 
cles when  the  other  group  is  made  to  contract. 
This  "reciprocal  innervation  of  antagonistic  mus- 
cles," as  Sherrington  has  called  it,^  is  thus  a 
device  for  orderly  action  in  the  body.  As  the 
above  description  has  shown,  there  are  peripheral 
oppositions  in  the  viscera  corresponding  to  the 
oppositions  between  flexor  and  extensor  muscles. 
In  all  probability  these  opposed  innervations  of 
the  viscera  have  counterparts  in  the  organization 
of  neurones  in  the  central  nervous  system.  Sher- 
rington has  noticed,  and  I  can  confirm  the  obser- 
vation, that  even  though  the  sympathetic  supply 
to  the  eye  is  severed  and  is  therefore  incapable  of 
causing  dilation  of  the  pupil,  nevertheless  the 
pupil  dilates  in  a  paroxysm  of  anger — due,  no 
doubt  (because  the  response  is  too  rapid  to  be 
mediated  by  the  blood  stream),  to  central  inhibi- 
tion of  the  cranial  nerve  supply  to  the  constrictor 
muscles — i.  e.,  an  inhibition  of  the  muscles  which 
naturally  oppose  the  dilator  action  of  the  sym- 
pathetic.    Pain,   the  major  emotions — fear  and 

duce  increased  tone  and  contraction  in  these  organs.     This 
double  action  of  the  vagus  is  marked  thus,  h=  ,  in  Tig.  1. 


36  BODILY    CHANGES 

rage — and  also  intense  excitement,  are  manifested 
in  the  activities  of  the  sympathetic  division. 
When  in  these  states  impulses  rush  out  over  the 
neurones  of  this  division  they  produce  all  the 
changes  typical  of  sympathetic  excitation,  such 
as  dilating  the  pupils,  inhibiting  digestion,  caus- 
ing pallor,  accelerating  the  heart,  and  various 
other  well-known  effects.  The  impulses  of  the 
sympathetic  neurones,  as  indicated  by  their  domi- 
nance over  the  digestive  process,  are  capable  of 
readily  overwhelming  the  conditions  established 
by  neurones  of  the  cranial  division  of  the  auto- 
nomic system. 

Neurones  of  the  Sympathetic  Division  and  Adrenal 
Secretion  Have  the  Same  Action 

Lying  anterior  to  each  kidney  is  a  small  body — 
the  adrenal  gland.  It  is  composed  of  an  external 
portion  or  cortex,  and  a  central  portion  or  me- 
dulla. From  the  medulla  can  be  extracted  a  sub- 
stance, called  variously  suprarenin,  adrenin,  epi- 
nephrin  or  "adrenalin,"*  which,  in  extraordinarily 
minute  amounts,  affects  the  structures  innervated 
by  the  sympathetic  division  of  the  autonomic  sys- 

*  The  name  "adrenalin"  is  proprietary.  "Epinephrin"  and 
"adrenin"  have  been  suggested  as  terms  free  from  commer- 
cial suggestions.  As  adrenin  is  shorter  and  more  clearly 
related  to  the  common  adjectival  form,  adrenal,  I  have  fol- 
lowed Schafer  in  using  adrenin  to  designate  the  substance 
produced  physiologically  by  the  adrenal  glands. 


VISCERAL    NEEVES  37 

tern  precisely  as  if  they  were  receiving  nervous 
impulses.  For  example,  when  adrenin  is  injected 
into  the  blood,  it  will  cause  pupils  to  dilate,  hairs 
to  stand  erect,  blood  vessels  to  be  constricted,  the 
activities  of  the  alimentary  canal  to  be  inhibited, 
and  sugar  to  be  liberated  from  the  liver.  These 
effects  are  not  produced  by  action  of  the  substance 
on  the  central  nervous  system,  but  by  direct  ac- 
tion on  the  organ  itself.'^  And  the  effects  oc- 
cur even  after  the  structures  have  been  removed 
from  the  body  and  kept  alive  artificially. 

The  adrenals  are  glands  of  internal  secretion, 
i.  e.,  like  the  thyroid,  parathyroid,  and  pituitary 
glands,  for  example ;  they  have  no  connection  with 
the  surface  of  the  body,  and  they  give  out  into 
the  blood  the  material  which  they  elaborate.  The 
blood  is  carried  away  from  each  of  them  by  the 
lumbo-adrenal  vein  which  empties  either  into  the 
renal  vein  or  directly  into  the  inferior  vena  cava 
just  anterior  to  the  openings  of  the  renal  veins. 
The  adrenal  glands  are  supplied  by  preganglionic 
fibres  of  the  autonomic  group,^  shown  in  solid 
line  in  Fig.  1.  This  seems  an  exception  to  the 
general  rule  that  gland  cells  have  an  outlying 
neurone  between  them  and  the  neurones  of  tho 
central  nervous  system.  The  medulla  of  the  adre- 
nal gland,  however,  is  composed  of  modified  nerve 
cells,  and  may  therefore  be  regarded  as  offering 
exceptional  conditions. 


38  BODILY    CHANGES 

The  foregoing  brief  sketch  of  the  organization 
of  the  autonomic  system  brings  out  a  number  of 
points  that  should  be  of  importance  as  bearing 
on  the  nature  of  the  emotions  which  manifest 
themselves  in  the  operations  of  this  system.  Thus 
it  is  highly  probable  that  the  sympathetic  division, 
because  arranged  for  diffuse  discharge,  is  likely 
to  be  brought  into  activity  as  a  whole,  whereas 
the  sacral  and  cranial  divisions,  arranged  for 
particular  action  on  separate  organs,  may  operate 
in  parts.  Also,  because  antagonisms  exist  be- 
tween the  middle  and  either  end  division  of  the 
autonomic,  affective  states  may  be  classified  ac- 
cording to  their  expression  in  the  middle  or  an 
end  division  and  these  states  would  be,  like  the 
nerves,  antagonistic  in  character.  And  finally, 
since  the  adrenal  glands  are  innervated  by  au- 
tonomic fibres  of  the  mid-division,  and  since  ad- 
renal secretion  stimulates  the  same  activities  that 
are  stimulated  nervously  by  this  division,  it  is 
possible  that  disturbances  in  the  realm  of  the 
sympathetic,  although  initiated  by  nervous  dis- 
charge, are  .automatically  augmented  and  pro- 
longed through  chemical  effects  of  the  adrenal 
secretion. 

EEFEKENCES 

^  Cannon :  The  American  Journal  of  Psychology,  1914, 
XXV,  p.  257. 


VISCERAL   NERVES  39 

-  For  a  summary  of  his  studies  of  the  organization  of  the 
autonomic  system,  see  Langley:  Ergebnisse  der  Physiologic, 
Wiesbaden,  1903,  ii^,  p.  818. 

^  See  Cannon :  American  Journal  of  Physiology,  1905, 
xiii,  p.  xxii. 

*  See  Sherrington :  The  Integrative  Action  of  the  Nerv- 
ous System,  New  York,  1909,  p.  19. 

^  Langley  and  Anderson :  Journal  of  Physiology,  1895, 
xix,  see  pp.  85,  122. 

^Sherrington:  Loc.  cit.,  p.  90. 

''  Elliott :  Journal  of  Physiology,  1905,  xxxii,  p.  426. 

8  See  Elliott :  Journal  of  Physiology,  1913,  xlvi,  p.  289  ff. 


CHAPTEE   III 

METHODS    OF   DEMONSTEATING   ADEENAL 
SECEETION  AND    ITS    NEEVOUS    CONTEOL 

As  stated  in  the  first  chapter,  the  inhibition  of 
gastric  secretion  produced  by  great  excitement 
long  outlasts  the  presence  of  the  object  which 
evokes  the  excitement.  The  dog  that  was  en- 
raged by  seeing  a  cat  for  five  minutes  secreted 
only  a  few  drops  of  gastric  juice  during  the  next 
fifteen  minutes.  Why  did  the  state  of  excitation 
persist  so  long  after  the  period  of  stimulation  had 
ended?  This  question,  which  presented  itself  to 
me  while  reading  Bickel  and  Sasaki's  paper,  fur- 
nished the  suggestion  expressed  at  the  close  of 
the  last  chapter,  that  the  excitement  might  pro- 
voke a  flow  of  adrenal  secretion,  and  that  the 
changes  originally  induced  in  the  digestive  organs 
by  nervous  impulses  might  be  continued  by  circu- 
lating adrenin.  The  prolongation  of  the  effect 
might  be  thus  explained.  Whether  that  idea  is 
correct  or  not  has  not  been  tested.  Its  chief  serv- 
ice was  in  leading  to  an  enquiry  as  to  whether 

40 


ADRENAL  SECRETION  41 

the  adrenal  glands  are  in  fact  stimulated  to  action 
in  emotional  excitement.  The  preganglionic  fibres 
passing  to  the  glands  are  contained  in  the  splanch- 
nic nerves.  What  is  the  effect  of  splanchnic  stim- 
ulation 1 

The  Evidence  that  Splanchnic  Stimulation  Induces 
Adrenal  Secretion 

It  was  in  1891  that  Jacobi  ^  described  ner^^e 
fibres  derived  from  the  splanchnic  trunks  which 
were  distributed  to  the  adrenal  glands.  Six  years 
later  Biedl  -  found  that  these  nerves  conveyed 
vaso-dilator  impulses  to  the  glands,  and  he  sug- 
gested that  they  probably  conveyed  also  secre- 
tory imjDulses.  Evidence  in  support  of  this  sug- 
gestion was  presented  the  following  year  by 
Dreyer,^  who  demonstrated  that  electrical  ex- 
citation of  the  splanchnic  nerves  produced  in  the 
blood  taken  from  the  adrenal  veins  an  increased 
amount  of  a  substance  having  the  power  of  rais- 
ing arterial  blood  pressure,  and  that  this  result 
was  independent  of  accompanying  changes  in  the 
blood  supply  to  the  glands.  The  conclusion  drawn 
by  Dreyer  that  this  substance  was  adrenin  has 
been  confirmed  in  various  ways  by  later  observers. 
Tscheboksaroff  ^  repeated  Dreyer's  procedure 
and  found  in  blood  taken  from  the  veins  after 
splanchnic  stimulation  evidences  of  the  presence 
of  adrenin  that  were  previously  absent.    Asher  ° 


42  BODILY   CHANGES 

observed  a  rise  of  blood  pressure  when  the  glands 
were  stimulated  in  such  a  manner  as  not  to  cause 
constriction  of  the  arteries — the  rise  was  there- 
fore assumed  to  be  due  to  secreted  ad*renin. 
Dilation  of  the  pupil  was  used  by  Meltzer  and 
Joseph  ^  to  prove  secretory  action  of  the  splanch- 
nics  on  the  adrenal  glands ;  they  found  that  stim- 
ulation of  the  distal  portion  of  the  cut  splanchnic 
nerve  caused  the  pupil  to  enlarge — an  effect  char- 
acteristic of  adrenin  circulating  in  the  blood. 
Elliott '^  repeated  this  procedure,  but  made  it 
a  more  rigorous  proof  of  internal  secretion  of  the 
adrenals  by  noting  that  the  effect  failed  to  ap- 
pear if  the  gland  on  the  stimulated  side  was  re- 
moved. Additional  proof  was  brought  by  myself 
and  Lyman  ^  when  we  found  that  the  typical 
drop  in  arterial  pressure  produced  in  cats  by  in- 
jecting small  amounts  of  adrenin  could  be  ex- 
actly reproduced  by  stimulating  the  splanchnic 
nerves  after  the  abdominal  blood  vessels,  which 
contract  when  these  nerves  are  excited,  were  tied 
so  that  no  changes  in  them  could  occur  to  in- 
fluence the  rest  of  the  circulation. 

The  problem  of  splanchnic  influence  on  the  ad- 
renal glands  Elliott  attacked  by  a  still  different 
method.  Using,  as  a  measure,  the  graded  effects 
of  graded  amounts  of  adrenin  on  blood  pressure, 
he  was  able  to  assay  the  quantity  of  adrenin  in 
adrenal  glands  after  various  conditions  had  been 


ADRENAL  SECRETION  43 

allowed  to  prevail.  The  tests  were  made  on  eats. 
In  these  animals  each  adrenal  gland  is  supplied 
only  by  the  splanchnic  fibres  of  its  own  side,  and 
the  two  glands  normally  contain  almost  exactly  the 
sarne  amomit  of  adrenin.  Elliott  ^  found  that 
when  the  gland  on  one  side  was  isolated  by  cutting 
its  splanchnic  supply,  and  then  impulses  were  sent 
along  the  intact  nerves  of  the  other  side,  either 
by  disturbing  the  animal  or  by  artificial  excita- 
tion of  the  nerves,  the  gland  to  which  these  fibres 
reached  invariably  contained  less  adrenin,  often 
very  much  less,  than  the  isolated  gland.  Results 
obtained  by  the  method  employed  by  Elliott  have 
been  confirmed  with  remarkable  exactness  in  re- 
sults obtained  by  Folin,  Denis  and  myself,^^ 
using  a  highly  sensitive  color  test  after  adding 
the  gland  extract  to  a  solution  of  phosphotungstic 
acid. 

All  these  observations,  with  a  variety  of  meth- 
ods, and  by  a  respectable  number  of  reliable  in- 
vestigators, are  harmonious  in  bringing  proof 
that  artificial  stimulation  of  the  nerves  leading 
to  the  adrenal  glands  will  induce  secretory  ac- 
tivity in  the  adrenal  medulla,  and  that  in  conse- 
quence adrenin  will  be  increased  in  the  blood. 
The  fact  is  therefore  securely  established  that  in 
the  body  a  mechanism  exists  by  which  these 
glands  can  be  made  to  discharge  this  peculiar  sub- 
stance promptly  into  the  circulation. 


44  BODILY   CHANGES 

The  Question  of  Adrenal  Secretion  in  Emotional 
Excitement 

As  we  have  already  seen,  the  phenomena  of  a 
great  emotional  disturbance  in  an  animal  indi- 
cate that  sympathetic  impulses  dominate  the  vis- 
cera. When,  for  example,  a  cat  becomes  fright- 
ened, the  pupils  dilate,  the  activities  of  the 
stomach  and  intestines  are  inhibited,  the  heart 
beats  rapidly,  the  hairs  of  the  back  and  tail  stand 
erect — from  one  end  of  the  animal  to  the  other 
there  are  abundant  signs  of  nervous  discharges 
along  sympathetic  courses.  Do  not  the  adrenal 
glands  share  in  this  widespread  subjugation  of 
the  viscera  to  sympathetic  control? 

This  question,  whether  the  common  excitements 
of  an  animal's  life  might  be  capable  of  evoking 
a  discharge  of  adrenin,  was  taken  up  by  D.  de  la 
Paz  and  myself  in  1910.  We  made  use  of  the  nat- 
ural enmity  between  two  laboratory  animals,  the 
dog  and  the  cat,  to  pursue  our  experiments.  In 
these  experiments  the  cat,  fastened  in  a  comfor- 
table holder  (the  holder  already  mentioned  as  be- 
ing used  in  X-ray  studies  of  the  movements  of 
the  alimentary  canal),  was  placed  near  a  barking 
dog.  Some  cats  when  thus  treated  showed  al- 
most no  signs  of  fear;  others,  with  scarcely  a 
movement  of  defence,  presented  the  typical  pic- 
ture. In  favorable  cases  the  excitement  was  al- 
lowed to  prevail  for  five  or  ten  minutes,  and  in 


ADRENAL  SECRETION  45 

a  few  cases  longer.    Samples  of  blood  were  taken 
within  a  few  minutes  before  and  after  the  period. 

The  Method  of  Securing  Blood  from  Near  the  Adrenal 

Veins 

The  blood  was  obtained  from  the  inferior  vena 
cava  anterior  to  the  opening  of  the  adrenal  veins, 
i.  e.,  at  a  point  inside  the  bodj^  near  the  level  of 
the  notch  at  the  lower  end  of  the  sternum.  To 
get  the  blood  so  far  from  the  surface  without 
disturbing  the  animal  was  at  first  a  difficult  prob- 
lem. We  found,  however,  that  by  making  anes- 
thetic with  ethyl  chloride  the  skin  directly  over 
the  femoral  vein  high  in  the  groin,  the  vein  could 
be  quickly  l)ared,  cleared  of  connective  tissue, 
tied,  and  opened  without  causing  any  general  dis- 
turbance whatever.  A  long,  fine,  flexible  catheter 
(2.4  millimeters  in  diameter)  which  had  pre- 
viously been  coated  with  vaseline  inside  and  out, 
to  lubricate  it  and  to  delay  the  clotting  of  blood 
within  it,  was  now  introduced  into  the  opening  in 
the  femoral  vein,  thence  through  the  iliac  and 
on  into  the  inferior  cava  to  a  point  near  the  level 
of  the  sternal  notch.  A  thread  tied  around  this 
tube  where,  after  being  inserted  to  the  proper  dis- 
tance, it  disappeared  into  the  femoral  vein, 
marked  the  extent  of  insertion,  and  permitted  a 
later  introduction  to  the  same  extent.  This  slight 
operation — a  venesection,  commonly  practised  on 


46  BODILY   CHANGES 

our  ancestors — consumed  only  a  few  minutes,  and 
as  the  only  possibility  of  causing  pain  was 
guarded  against  by  local  anesthesia,  the  animal 
remained  tranquil  throughout.  Occasionally  it 
was  necessary  to  stroke  the  cat's  head  gently  to 
keep  her  quiet  on  the  holder,  and  under  such  cir- 
cumstances I  have  known  her  to  purr  during  all 
the  preparations  for  obtaining  the  blood,  and 
while  the  blood  was  being  taken. 

The  blood  (3  or  4  cubic  centimetres)  was  slowly 
drawn  through  the  catheter  into  a  clean  glass 
syringe.  Care  was  taken  to  avoid  any  marked 
suction  such  as  might  cause  collapse  of  the  vein 
near  the  inner  opening  of  the  tube.  As  soon  as 
the  blood  was  secured,  the  catheter  was  removed 
and  the  vein  tied  loosely,  to  prevent  bleeding. 
The  blood  was  at  once  emptied  into  a  beaker,  and 
the  fibrin  whipped  from  it  by  means  of  fringed 
rubber  tubing  fitted  over  a  glass  rod.  Since  this 
defibrinated  blood  was  obtained  while  the  ani- 
mal was  undisturbed,  it  was  labelled  "quiet 
blood." 

The  animal  was  then  exposed  to  the  barking 
dog,  as  already  described,  and  immediately  there- 
after blood  was  again  removed,  from  precisely 
the  same  region  as  before.  This  sample,  after 
being  defibrinated,  was  labelled  "excited  blood." 
The  two  samples,  the  "quiet"  and  the  "excited," 
both  obtained  in  the   same  manner  and  subse- 


ADEEXAL  SECRETION  47 

quently  treated  in  the  same  manner,  Tvere  now 
tested  for  their  content  of  adrenin. 

The  ^Method  of  Testing  the  Blcxjo  for  Adrenix 

It  was  desirable  to  use  as  a  test  tissues  to 
which  the  blood  was  naturally  related.  As  will 
be  recalled,  adrenin  affects  viscera  even  after 
they  have  been  removed  from  the  body,  just  as  if 
they  were  receiving  impulses  via  sympathetic 
fibres,  and  further,  that  sympathetic  fibres  nor- 
mally deliver  impulses  which  cause  contraction 
of  the  internal  genitals  and  relaxation  of  the 
stomach  and  intestines.  The  uterus  has  long  been 
employed  as  a  test  for  adrenin,  the  presence  of 
which  it  indicates  by  increased  contraction.  That 
isolated  strips  of  the  longitudinal  muscle  of  the 
intestine,  which  are  contracting  rhythmically,  are 
characteristically  inhibited  by  adrenin  in  dilu- 
tions of  1  part  in  20  millions,  had  been  shown  by 
Magnus  in  1905.  Although,  previous  to  our  in- 
vestigation in  1910,  this  extremely  delicate  reac- 
tion had  not  been  used  as  a  biological  signal  for 
adrenin,  it  possesses  noteworthy  advantages  over 
other  methods.  The  intestine  is  found  in  all  ani- 
mals and  not  in  only  half  of  them,  as  is  the  uterus ; 
it  is  ready  for  the  test  within  a  few  minutes,  in- 
stead of  the  several  hours  said  to  be  required  for 
the  best  use  of  the  uterus  preparation;^^  and  it 
responds  by  relaxing.     This  last  characteristic 


48  BODILY   CHANGES 

is  especially  important,  for  in  defibrinated  blood 
there  are,  besides  adrenin,  other  substances  cap- 
able of  causing  contraction  of  smooth  muscle,^^ 
and  liable  therefore  to  lead  to  erroneous  con- 
clusions when  a  structure  which  responds  by  con- 
tracting, such  as  uterus  or  artery,  is  used  to  prove 
whether  adrenin  is  present.  On  the  other  hand, 
substances  producing  relaxation  of  smooth  muscle 
are  few,  and  are  unusual  in  blood.^^ 

We  used,  therefore,  the  strip  of  intestinal  mus- 
cle as  an  indicator.  Later  Ho  skins  ^^  modified 
our  procedure  by  taking,  instead  of  the  strip,  a 
short  segment  of  the  rabbit  intestine.  The  seg- 
ment is  not  subjected  to  danger  of  injury  during 
its  preparation,  and  when  fresh  it  is  almost  in- 
credibly sensitive.  It  may  be  noticeably  ihhibited 
by  adrenin,  1  part  in  200  millions! 

The  strip,  or  the  intestinal  segment,  was  sus- 
pended between  minute  wire  pincers  {serres 
fines)  in  a  cylindrical  chamber  8  millimeters  in 
diameter  and  5  centimeters  deep.  By  a  thread 
attached  to  the  lower  serre  fine  the  preparation 
was  drawn  into  the  chamber,  and  was  held  firmly; 
by  the  upper  one  it  was  attached  to  the  short  end 
of  a  writing  lever  (see  Fig.  2).  When  not  ex- 
posed to  blood,  the  strip  was  immersed  in  a  nor- 
mal solution  of  the  blood  salts  (Einger's).  The 
blood  or  the  salt  solution  could  be  quickly  with- 
drawn from  or  introduced  into  the  chamber,  with- 


ADRENAL  SECRETION 


49 


out  disturbing  the  muscle,  by  means  of  a  fine 
pipette  passed  do^^^l  along  the  inner  surface.  The 
chamber  and  its  contents,  the  stock  of  Ringer's 


Figure  2. — Diagram  of  the  arrangements  for  recording  con- 
tractions of  the  intestinal  muscle. 


solution,  and  the  samples  of  "quiet"  and  "ex- 
cited" blood  were  all  surrounded  by  a  large  vol- 
ume of  water  kept  approximately  at  body  tem- 
perature (37°  C).  Through  the  blood  or  the  salt 
solution  in  the  chamber  oxygen  was  passed  in  a 
slow  but  steady  stream  of  bubbles.  Under  these 
circumstances  the  strip  will  live  for  hours,  and 
will  contract  and  relax  in  a  beautifully  regular 
rhythm,  which  may  be  recorded  graphically  by 
the  writing  lever. 

The  first  effect  of  surrounding  the  muscle  with 
blood,  whether  "quiet"  or  "excited,"  was  to  send 
it  into  a  strong  contraction  which  might  persist, 
sometimes  with  slight  oscillations,  for  a  minute 
or  two  (see  Figs.  4  and  5).  After  the  initial  short- 
ening, the  strip,  if  in  quiet  ))lood  soon  began  to 


50  BODILY    CHANGES 

contract  and  relax  rhythmically  and  with  each  re- 
laxation to  lengthen  more,  nntil  a  fairly  even 
base  line  appeared  in  the  written  record.  At  this 
stage  the  addition  of  fresh  "quiet"  blood  usually 
had  no  effect,  even  though  the  strip  were  washed 
once  with  Einger's  solution  before  the  second  por- 
tion of  the  blood  was  added.  For  comparison  of 
the  effects  of  "quiet"  and  "excited"  blood  on  the 
contracting  strip,  the  two  samples  were  each 
added  to  the  muscle  immediately  after  the  Eing- 
er's solution  had  been  removed,  or  they  were  ap- 
plied to  the  muscle  alternately  and  the  differences 
in  effect  then  noted.  The  results  obtained  by 
these  methods  are  next  to  be  presented. 

EEFEEENCES 

^  Jacobi :  Archiv  f iir  experimentelle  Patbologle  und  Phar- 
makologie,  1891,  xxix,  p.  185. 

2  Biedl :  Archiv  f iir  die  gesammte  Physiologie,  1897,  Ixvii, 
pp.  456,  481. 

^  Dreyer :  American  Journal  of  Physiology,  1898-99,  ii, 
p.  219. 

*  Tscheboksaroff :  Archiv  f  iir  die  gesammte  Physiologie, 
1910,  cxxxvii,  p.  103. 

^  Asher :  Zeitschrift  fiir  Biologie,  1912,  Iviii,  p.  274. 

^  Meltzer  and   Joseph :   American   Journal   of  Physiology, 

1912,  xxix,  p.  xxxiv. 

^Elliott:  Journal  of  Physiology,  1912,  xliv,  p.  400. 

^  Cannon  and  Lyman :  American  Journal  of  Physiology, 

1913,  xxxi,  p.  377. 

9  Elliott :  Journal  of  Physiology,  1912,  xliv,  p.  400. 
^°  Folin,  Cannon  and  Denis :  Journal  of  Biological  Chem- 
istry, 1913,  xiii,  p.  477. 


ADRENAL  SECRETION  51 

^^  Fraenkel :  Arcliiv  fiir  experimentelle  Pathologie  und 
Pharmakologie,  1909,  Ix,  p.  399. 

^-  See  O'Connor :  Archiv  fiir  die  experimentelle  Patholo- 
gie und  Pharmakologie,  1912,  Ixvii,  p.  206. 

^2  Grutzner :  Ergebnisse  der  Physiologic,  1904,  iii^,  p.  66 ; 
Magnus :  Loc.  cit.,  p.  69. 

i-'Hoskins:  Journal  of  Pharmacology  and  Experimental 
Therapeutics,  1911,  iii,  p.  95. 


CHAPTEE   IV 

ADEENAL   SECEETION  IN   STRONG  EMOTIONS 
AND   PAIN 

If  the  secretion  of  adrenin  is  increased  in  strong 
emotional  states  and  in  pain,  that  constitutes  a 
fact  of  considerable  significance,  for,  as  already 
mentioned,  adrenin  is  capable  of  producing  many 
of  the  bodily  changes  which  are  characteristically 
manifested  in  emotional  and  painful  experiences. 
It  is  a  matter  of  prime  importance  for  further 
discussion  to  determine  whether  the  adrenal 
glands  are  in  fact  roused  to  special  activity  in 
times  of  stress. 

The  Evidence  that  Adrenal  Secretion  Is  Increased  in 
Emotional  Excitement 

That  blood  from  the  adrenal  veins  causes  the 
relaxation  of  intestinal  muscle  characteristic  of 
adrenal  extract  or  adrenin  is  shown  in  Fig.  3. 
The  muscle  was  originally  beating  in  blood  which 
contained  no  demonstrable  amount  of  adrenal  se- 
cretion ;  this  inactive  blood  was  replaced  by  blood 

52 


ADRENAL   SECRETION   IN  EMOTIONS     53 

from  the  adrenal  veins,  obtained  after  quick 
etherization.  Etherization,  it  will  be  recalled,  is 
accompanied  by  a  "stage  of  excitement."  Re- 
laxation occurred  almost  immediately  (at  h). 
Then   the   rhythm   was   renewed   in   the   former 


Figure  3. — Intestinal  muscle  beat- 
ing in  inactive  blood,  which  was  with- 
drawn from  the  chamber  at  a.  Blood 
from  the  adrenal  vein  of  an  animal  ex- 
cited by  etherization  was  substituted 
at  h,  and  withdrawn  at  c.  Contrac- 
tions were  restored  in  the  original  in- 
active blood  which  was  removed  at  d. 
Blood  from  the  renal  vein  (same  ani- 
mal) was  added  at  e. 

In  this  and  subsequent  records  time 
is  marked  in  half  minutes. 

blood,  and  thereupon  the  muscle  was  surrounded 
with  blood  from  the  vein  leading  away  from  the 
left  kidney,  i.  e.,  blood  obtained  from  the  same 
animal  and  under  the  same  conditions  as  the 
adrenal  blood,  but  from  a  neighboring  vein.  No 
relaxation  occurred.  By  this  and  other  similar 
tests  the  reliability  of  the  method  was  proved. 


64  BODILY    CHANGES 

In  no  instance  did  blood  from  the  inferior  vena 
cava  of  the  quiet  normal  animal  produce  relaxa- 
tion. On  the  other  hand,  blood  from  the  animal 
after  emotional  excitement  showed  more  or  less 
promptly  the  typical  relaxation.     In  Fig.  4  is 


Figure  4. — Alternate  application  of  "excited"  blood  (at  6 
and/)  and  "quiet"  blood  (at  d),  from  the  same  animal,  to  in- 
testinal muscle  initially  beating  in  Ringer's  solution. 


represented  the  record  of  intestinal  muscle  which 
was  beating  regularly  in  Ringer's  solution.  At  a 
the  Einger's  solution  was  removed,  and  at  h  "ex- 
cited" blood  was  added;  after  the  preliminary 
shortening,  which,  as  already  stated,  occurs  at 
the  first  immersion  in  blood,  the  muscle  length- 
ened gradually  into  complete  inhibition.  At  c  the 
"excited"  blood  was  removed,  and  at  d  "quiet" 
blood  was  added  in  its  place.  The  muscle  at  once 
began  fairly  regular  rhythmic  beats.  At  e  the 
"quiet"  blood  was  removed,  and  at  /  the  "excited" 
blood  was  again  applied.  The  muscle  lengthened 
almost  immediately  into  an  inhibited  state.  In 
this  instance  the  "excited"  blood  was  taken  after 


ADRENAL   SECRETION  IN  EMOTIONS     55 

the  cat  had  been  barked  at  for  about  fifteen  min- 
utes. 

The  increase  of  effect  with  prolongation  of  the 
period  of  excitement  is  shown  in  Fig.  5.    A  is  the 


Figure  5. — The  effect  of  prolonging  the  excitement.  A,  the 
record  in  "quiet"  serum;  B,  in  defibrinated  blood  after  eleven 
minutes  of  excitement;  and  C,  in  serum  after  fifteen  minutes  of 
excitement. 

record  of  contractions  after  the  muscle  was  sur- 
rounded with  "quiet"  blood  serum.  B  shows  the 
gradual  inhibition  which  occurred  when  the  mus- 
cle was  surrounded  with  defibrinated  blood  taken 
when  the  animal  had  been  excited  eleven  minutes. 
And  C  is  the  record  of  rapid  inhibition  after  fif- 
teen minutes  of  excitement.  In  other  instances 
the  effect  was  manifested  merely  by  a  lowering 
of  the  tonus  of  the  muscle,  and  a  notable  slowing 
of  the  beats,  without,  however,  a  total  abolition 
of  them. 

The  inference  that  this  inhibition  of  contrac- 
tion of  the  intestinal  muscle  is  due  to  an  increased 
amount   of    adrenal    secretion   in   the   "excited'* 


56 


BODILY   CHANGES 


blood  de  la  Paz  and  I  justified  on  several  grounds : 
(1)  The  inhibition  was  produced  by  "excited" 
blood  from  the  inferior  vena  cava  anterior  to  the 
mouths  of  the  adrenal  veins,  when  blood  from  the 
femoral  vein,  taken  at  the  same  time,  had  no  in- 
hibitory influence.  Since  blood  from  the  femoral 
vein  is  typical  of  the  cava  blood  below  the  en- 
trance of  the  kidney  veijis,  the  conclusion  is  war- 
ranted that  the  difference  of  effect  of  the  two 
samples  of  blood  is  not  due  to  any  agent  below 
the  kidneys.  But  that  blood  from  the  kidneys 
does  not  cause  the  relaxation  is  shown  in  Fig.  3. 


FiGXJEE  6. — Failure  of  the  cava  blood 
(added  at  a)  to  produce  inhibition  when 
excitement  has  occurred  after  removal 
of  the  adrenal  glands.  The  muscle  later 
proved  sensitive  to  adrenin  in  blood  in 
the   ratio    1:1,000,000. 

The  only  other  structures  which  could  alter  the 
blood  between  the  two  points  at  which  it  was 
taken  are  the  adrenal  glands,  and  the  material 


ADEENAL  SECRETION  IN  EMOTIONS     57 

secreted  by  them  would  produce  precisely  the 
inhibition  of  contraction  which  was  in  fact  pro- 
duced. 

(2)  If  in  ether  anesthesia  the  blood  vessels 
leading  to  and  from  the  adrenal  glands  are  first 
carefully  tied,  and  then  the  glands  are  removed,  ex- 


FiGURE  7.— Effect  of  adding  adrenin  1:1,000,000  (A),  1:2,000,000 
(B),  and  1:3,000,000  (Cj,  to  formerly  inactive  blood.  In  each  case 
a  marks  the  moment  when  the  quiet  blood  %vaa  removed,  and  b,  the 
time  when  the  blood  with  adrenin  was  added. 


citement  four  or  five  hours  later,  before  the  weak- 
ness that  follows  the  removal  has  become  promi- 
nent, does  not  alter  the  blood  so  that  the  typical 
inhibition  occurs  (see  Fig.  6).  Thus,  although 
the  animal  shows  all  the  characteristic  signs  of 
sympathetic  stimulation,  the  blood,  in  the  absence 
of  the  adrenals,  remains  unchanged. 

(3)  As  already  shown,  sometimes  the  effect  pro- 


58  BODILY    CHANGES 

duced  by  the  "excited"  blood  was  prompt  inbibi- 
tion,  sometiines  the  inhibition  followed  only  after 
several  beats,  and  sometimes  a  slowing  and  short- 
ening of  contractions,  with  a  lower  tone,  were  the 
sole  signs  of  the  action  of  adrenin.  All  these 
degrees  of  relaxation  can  be  duplicated  by  adding 
to  inactive  blood  varying  amounts  of  adrenin. 
Fig.  7  shows  the  effects,  on  a  somewhat  insensi- 
tive muscle  preparation,  of  adding  adrenin, 
1:1,000,000  (A),  1:2,000,000  (B),  and  1:3,000,000 
( C ) ,  to  different  samples  of  blood  previously  with- 
out inhibitory  influence.  These  effects  of  adrenin 
and  the  effects  jDroduced  by  blood  taken  near  the 
opening  of  the  adrenal  veins  are  strikingly  analo- 
gous. 

(4)  Emden  and  v.  Furth^  have  reported  that 
0.1  gram  of  suprarenin  chloride  disappears  almost 
completely  in  two  hours  if  added  to  200  cubic 
centimeters  of  defibrinated  beef  blood,  and  the 
mixture  constantly  aerated  at  body  temjoerature. 
"Excited"  blood  which  produces  inhibition  loses 
that  power  on  standing  in  the  cold  for  twenty-four 
hours,  or  on  being  kept  warm  and  agitated  with 
bubbling  oxygen.  This  change  is  illustrated  in 
Fig.  8 ;  the  power  of  the  "excited"  blood  to  inhibit 
the  contractions  of  the  intestinal  muscle  when  rec- 
ord A  was  written  was  destroyed  after  three 
hours  of  exjDOSure  to  bubbling  oxygen,  as  shown 
by  record  B.     The  destruction   of  adrenin  and 


ADEENAL   SECRETION  IN  "EMOTIONS     59 

the   disappearance   of  the   effect  which  adrenin 
would  produce  are  thus  closely  parallel. 

All  these  considerations,  taken  with  the  proof 


Figure  8. — The  effect  of  bubbling 
oxygen  through  active  blood.  A,  re- 
laxation after  active  blood  applied  at 
a;  B,  failure  of  relaxation  when  the 
same  blood,  oxygenated  three  hours, 
was  applied  to  a  fresh  strip  at  b. 

that  sympathetic  impulses  increase  secretion  of  the 
adrenal  glands,  and  taken  also  with  the  evidence 
that,  during  such  emotional  excitement  as  was  em- 
ployed in  these  experiments,  signs  of  sympathetic 
discharges  appeared  throughout  the  animal  from 
the  dilated  pupil  of  the  eye  to  the  standing  hairs 
of  the  tail-tip,  led  us  to  the  conclusions  that  the 
characteristic  action  of  adrenin  on  intestinal  mus- 
cle was  in  fact,  in  our  experiments,  due  to  secre- 
tion of  the  adrenal  glands,  and  that  that  secretion 
is  increased  in  great  emotion. 

The   Evidence  that   Adrenal   Secretion   is   Increased  by 
"Painful"  Stimulation 
As  mentioned  in  the  first  chapter,  stimulation  of 
sensory  fibres  in  one  of  the  larger  nerve  trunks 


60  BODILY    CHANGES 

is  known  to  result  in  such  nervous  discliarges  along 
sympathetic  paths  as  to  produce  marked  inhibi- 
tion of  digestive  processes.  Other  manifestations 
of  sympathetic  innervations — e.  g.,  contraction  of 
arterioles,  dilation  of  pupils,  erection  of  hairs — 
are  also  demonstrable.  And  since  the  adrenal 
glands  are  stimulated  to  activity  by  sympathetic 
impulses,  it  was  possible  that  they  would  be  af- 
fected as  are  other  structures  supplied  with  sym- 
pathetic fibres,  and  that  they  would  secrete  in 
greater  abundance  when  sensory  nerves  were  irri- 
tated. 

The  testing  of  this  possibility  was  undertaken  by 
Hoskins  and  myself  in  1911.  Since  bodily  changes 
from  "painful"  stimulation  can  in  large  degree  be 
produced  in  an  anesthetized  animal,  without,  how- 
ever, an  experience  of  pain  by  the  animal,  it  was 
possible  to  make  the  test  quite  simply.  The  sen- 
sory stimulus  was  a  rapidly  interrupted  induced 
current  applied  to  the  sciatic  nerve.  The  current 
was  increased  in  strength  as  time  passed,  and  thus 
the  intensity  of  the  effect,  indicated  by  continuous 
dilation  of  the  pupils,  was  maintained.  There  was 
no  doubt  that  such  stimulation  would  have  caused 
very  severe  pain  if  the  animal  had  not  been  anes- 
thetized. Indeed,  the  stimulus  used  was  probably 
much  stronger  than  would  be  necessary  to  obtain 
a  positive  result  in  the  absence  of  the  anesthetic 
(urethane),  which  markedly  lessens  the  irritabil- 


ADRENAL  SECRETION  IN  EMOTIONS     61 

ity   of   visceral   nerve   fibres.^     In   different  in- 
stances the  stimulation  lasted  from  three  to  six 
minutes.    Throughout  the  period  there  was  mark- 
edly increased  rapidity  and  depth  of  breathing. 
As  Fig.  9  shows,  the  normal  blood,  removed 


Figure  9. — Intestinal  mus- 
cle beating  in  normal  vena  cava 
blood,  removed  at  1  and  re- 
newed at  2.  At  3  normal  blood 
removed.  At  4  contraction  in- 
hibited by  vena  cava  blood 
drawn  after  sensory  stimula- 
tion; at  .5  removed.  At  6  Rin- 
ger's solution  substituted. 


62  BODILY    CHANGES 

from  the  vena  cava  before  stimulation,  caused  no 
inhibition  of  the  beating  segment,  whereas  that 
removed  afterwards  produced  a  deep  relaxation. 
Hoskins  and  I  showed  that  the  increased  respira- 
tion which  accompanies  "painful"  stimulation  does 
not  augment  adrenal  activity.  We  concluded, 
therefore,  that  when  a  sensory  trunk  is  strongly 
excited  the  adrenal  glands  are  reflexly  stimulated, 
and  that  they  pour  into  the  blood  stream  an  in- 
creased amount  of  adrenin. 

Confirmation  of  Our  Results  by  Other  Observers 

The  foregoing  experiments  and  conclusions  were 
reported  in  1911.  In  1912,  Elliott  ^  brought  con- 
firmatory evidence  by  use  of  a  method  quite  differ- 
ent from  ours.  As  previously  stated,  he  studied 
the  effects  of  experimental  procedures  on  adrenal 
secretion  by  a  careful  comparative  quantitative 
assay  of  the  adrenin  content  of  the  glands  when 
one  gland  was  isolated  from  the  central  nervous 
system  and  the  other  left  connected.  He  took 
advantage  of  the  action  of  morphia  and  of  the 
substance  B-tetrahydronaphthylamine  in  evoking 
in  cats  all  the  appearances  of  great  fright.  After 
the  animals  had  thus  been  "frightened,"  he  found 
that  the  adrenal  gland  which  was  still  connected 
with  the  spinal  cord  was  much  depleted  of  its 
adrenin  content  compared  with  the  other,  isolated 
gland.     And  he  observed,  further,  that  animals 


ADKENAL   SECRETION  IN  EMOTIONS     63 

newly  brought  to  the  laboratory,  and  evidently 
disturbed  by  the  strangeness  of  their  surroundings, 
had  a  considerably  smaller  amount  of  adrenin  in 
their  glands  than  other  animals  grown  accustomed 
to  the  situation.  Elliott  also  observed  that  joro- 
longed  excitation  of  a  sensory  nerve,  such  as  the 
great  sciatic,  may  cause  the  adrenin  largely  to 
disappear  from  the  gland  still  connected  with  the 
central  nervous  system  and  subjected,  therefore,  to 
reflex  influences. 

Our  conclusions  have  also  been  confirmed  more 
recently  (1913)  by  Hitchings,  Sloan  and  Austin,^ 
working  in  Crile's  laboratory  in  Cleveland.  They 
used  the  same  method  which  we  had  used  to  ob- 
tain blood  and  to  test  for  adrenin,  and  found  that 
after  great  fear  and  rage  had  been  induced  in  a 
cat  by  the  attempt  of  a  muzzled  dog  to  fight  it, 
the  adrenin  reaction  was  clearly  demonstrable. 
And  just  as  we  had  noted  that  the  reaction  did  not 
occur  if  the  adrenal  glands  had  been  removed,  they 
showed  that  it  did  not  occur  if  the  nervous  connec- 
tions with  the  spinal  cord  were  previously  severed. 

The  logic  of  all  these  experiments  may  be  briefly 
summed  up.  That  the  adrenal  glands  are  subject 
to  splanchnic  influence  has  been  demonstrated 
anatomically  and  by  the  physiological  effects  of 
their  secretion  after  artificial  stimulation  of  the 
splanchnic  nerves.  Impulses  are  normally  sent 
along  these  nerves,  in  the  natural  conditions  of 


64  BODILY    CHANGES 

life,  when  animals  become  greatly  excited,  as  in 
fear  and  rage  and  pain.  There  is  every  probabil- 
ity, therefore,  that  these  glands  are  stimulated  to 
extra  secretion  at  such  times.  Both  by  an  ex- 
ceedingly delicate  biological  test  (intestinal  mus- 
cle)  and  by  an  examination  of  the  glands  them- 
selves, clear  evidence  has  been  secured  that  in  pain 
and  deep  emotion  the  glands  do,  in  fact,  pour  out 
an  excess  of  adrenin  into  the  circulating  blood. 

Here,  then,  is  a  remarkable  group  of  phenomena 
— a  pair  of  glands  stimulated  to  activity  in  times 
of  strong  excitement  and  by  such  nerve  impulses 
as  themselves  produce  at  such  times  profound 
changes  in  the  viscera ;  and  a  secretion  given  forth 
into  the  blood  stream  by  these  glands,  which  is 
capable  of  inducing  by  itself,  or  of  augmenting, 
the  nervous  influences  which  induce  the  very 
changes  in  the  viscera  which  accompany  suffering 
and  the  major  emotions.  What  may  be  the  sig- 
nificance of  these  changes,  occurring  when  condi- 
tions of  pain  and  great  excitement — experiences 
common  to  animals  of  most  diverse  types  and 
probably  known  to  their  ancestors  for  ages  past 
— lay  hold  of  the  bodily  functions  and  determine 
the  instinctive  responses  ? 

Certain  remarkable  effects  of  injecting  adrenin 
into  the  blood  have  for  many  years  been  more  or 
less  well  recognized.  For  example,  when  injected 
it  causes  liberation  of  sugar  from  the  liver  into 


ADRENAL   SECRETION   IN  EMOTIONS     65 

the  blood  stream.  It  relaxes  the  smooth  muscle 
of  the  bronchioles.  Some  old  experiments  indi- 
cated that  it  acts  as  an  antidote  for  muscular 
fatigue.  It  alters  the  distribution  of  the  blood  in 
the  body,  driving  it  from  the  abdominal  viscera 
into  the  heart,  lungs,  central  nervous  system  and 
limbs.  And  there  was  some  evidence  that  it  ren- 
ders more  rapid  the  coagulation  of  the  blood. 
There  may  be  other  activities  of  adrenin  not  yet 
discovered — it  may  co-operate  with  the  products 
of  other  glands  of  internal  secretion.  And  other 
glands  of  internal  secretion  may  be  stimulated  by 
sympathetic  impulses.  But  we  were  not  concerned 
with  these  possibilities.  We  wished  to  know 
whether  the  adrenin  jDOured  out  in  pain  and  emo- 
tional excitement  produced  or  helped  to  produce 
the  same  effects  that  follow  the  injection  of  adre- 
nin. Our  later  researches  were  concerned  with  an- 
swers to  this  question. 

EEFERENCES 

^  Embden  and  v.  Furth :  Hofmeister's  Beitrage  zur 
chemischen  Physiologie  und  Pathologic,  1904,  iv,  p.  423. 

2  Elliott:  Journal  of  Physiology,  1905,  xxxii,  p.  448. 

3  Elliott:  Journal  of  Physiology,  1912,  xliv,  p.  409. 

*  Hitchings,  Sloan  and  Austin :  Cleveland  Medical  Jour- 
nal, 1913,  xii,  p.  G86;  see  also  Crile  and  Lower:  Anoci-asso- 
ciation,  Philadelphia,  1914,  p.  5G. 


CHAPTER   V 

THE    INCEEASE    OF    BLOOD    SUGAR    IN    PAIN 
AND   GEE  AT   EMOTION 

Sugar  is  the  form  in  which  carbohydrate  mate- 
rial is  transported  in  organisms ;  starch  is  the  stor- 
age form.  In  the  bodies  of  animals  that  have 
been  well  fed  the  liver  contains  an  abundance  of 
glycogen  or  "animal  starch,"  which  may  be  called 
upon  in  times  of  need.  At  such  times  the  glycogen 
is  changed,  and  set  free  in  the  blood  as  sugar. 
Ordinarily  there  is  a  small  percentage  of  sugar 
in  the  blood — from  0.06  to  0.1  per  cent.  When 
only  this  small  amount  is  present  the  kidneys  are 
capable  of  preventing  its  escape  ir  any  noteworthy 
amount.  If  the  percentage  rises  to  the  neighbor- 
hood of  0.2-0.3  per  cent,  however,  the  sugar  passes 
the  obstacle  set  up  by  the  kidneys,  and  is  readily 
demonstrable  in  the  urine  by  ordinary  tests.  The 
condition  of  "glycosuria,"  therefore,  may  prop- 
erly be  considered,  in  certain  circumstances,  as 
evidence  of  increased  sugar  in  the  blood.  The  in- 
jection of  adrenin  can  liberate  sugar  from  the 

66 


INCREASE    OF    BLOOD    SUGAR        67 

liver  to  such  an  extent  that  glycosuria  results. 
Does  the  adrenal  secretion  discharged  in  pain  and 
strong  emotional  excitement  play  a  role  in  pro- 
ducing glycosuria  under  such  conditions? 

In  clinical  literature  scattered  suggestions  are 
to  be  found  that  conditions  giving  rise  to  emo- 
tional states  may  be  the  occasion  also  of  more  or 
less  jDcrmanent  glycosuria.  Qreat  grief  and  pro- 
longed anxiety  during  a  momentous  crisis  have 
been  regarded  as  causes  of  individual  instances 
of  diabetes,  and  anger  or  fright  has  been  followed 
by  an  increase  in  the  sugar  excreted  by  persons 
who  already  have  the  disease.  Kleen  ^  cites 
the  instance  of  a  Ge-rman  officer  whose  diabetes 
and  whose  Iron  Cross  for  valor  both  came  from 
a  stressful  experience  in  the  Franco-Prussian  War. 
The  onset  of  the  disease  in  a  man  directly  after 
his  wife  was  discovered  in  adultery  is  described 
by  Naunyn;^  and  this  author  also  mentions 
two  cases  in  his  own  practice — one  started  during 
the  bombardment  of  Strassburg  (1870),  the  other 
started  a  few  days  after  a  companion  had  shot 
himself.  In  cases  of  mental  disease,  also,  states 
of  depression  have  been  described  accompanied 
by  sugar  in  the  urine.  Schultze^  has  reported 
that  in  these  cases  the  amount  of  glycosuria  is  de- 
pendent on  the  degree  of  depression,  and  that  the 
greatest  excretion  of  sugar  occurs  in  the  fear- 
psychoses.    Raimann  ^  has  reported  that  in  both 


68  BODILY   CHANGES 

melancliolia  and  mania  the  assimilation  limit  of 
sugar  may  be  lowered.  Similar  results  in  the 
insane  have  recently  been  presented  by  Mita,^ 
and  by  Folin  and  Denis.*^  The  latter  investiga- 
tors found  glycosuria  in  12  per  cent  of  192  insane 
patients,  most  of  whom  suffered  from  depression, 
apprehension,  or  excitement.  And  Arndt  ^  has 
observed  glycosuria  appearing  and  disappearing 
as  alcoholic  delirium  appeared  and  disappeared  in 
his  patients. 

Although  clinical  evidence  thus  indicates  an 
emotional  origin  of  some  cases  of  diabetes  and 
glycosuria,  the  intricacies  of  existence  and  the 
complications  of  disease  in  human  beings  throw 
some  doubt  on  the  value  of  that  evidence.  Both 
Naunyn  ^  and  Hirschf eld,  although  mentioning 
instances  of  diabetes  apparently  due  to  an  emo- 
tional experience,  urge  a  skeptical  attitude  to- 
ward such  statements.  It  is  desirable,  therefore, 
that  the  question  of  an  emotional  glycosuria  be 
tested  under  simpler  and  more  controllable  con- 
ditions. "Emotional  glycosuria"  in  experimental 
animals  has  indeed  been  referred  to  by  Water- 
man and  Smit  ^  and  more  recently  by  Hender- 
son and  Underhill.^^  Both  these  references,  how- 
ever, are  based  on  the  work  of  Bohm  and  Hoff- 
mann,^^  reported  in  1878. 


INCREASE    OF    BLOOD    SUGAR        69 

Glycosuria  From  Pain 

Bolim  and  Hoffmann  found  that  cats,  when 
bound  to  an  operating  board,  a  tube  inserted  into 
the  trachea  (without  anesthesia),  and  in  some 
instances  a  catheter  inserted  into  the  urethra 
through  an  opening  above  the  pubis,  had  in  about 
half  an  hour  an  abundance  of  sugar  in  the  urine. 
In  three  determinations  sugar  in  the  blood  proved 
slightly  above  "normal"  so  long  as  sugar  was  ap- 
pearing in  the  urine,  but  returned  to  "normal" 
as  the  glycosuria  disappeared.  Since  they  were 
able  to  produce  the  phenomenon  by  simply  bind- 
ing animals  to  the  holder,  they  called  it  "Fes- 
selungsdiabetes." 

As  possible  causes  of  this  glycosuria  in  bound 
animals,  they  considered  opening  the  trachea, 
cooling,  and  pain.  The  first  two  they  readily 
eliminated,  and  still  they  found  sugar  excreted. 
Pain  they  could  not  obviate,  and  since,  without 
binding  the  animals,  they  caused  glycosuria  by 
merely  stimulating  the  sciatic  nerves,  they  con- 
cluded that  painful  confinement  was  itself  a  suffi- 
cient cause.  Other  factors,  however,  such  as  cool- 
ing and  circulatory  disturbances,  probably  co- 
operated with  pain,  they  believed,  to  produce  the 
result.  Their  observations  on  cats  have  been 
proved  true  also  of  rab])its;^-  and  recently  it  has 
been  shown  that  an  operation  involving  some  pain 
increases  blood  sugar  in  dogs.^^    Temporary  gly- 


70  BODILY    CHANGES 

cosnria  has  likewise  been  noted  in  association  with 
intense  pain  in  human  beings. 

Inasmuch  as  Bohm  and  Hoffmann  did  not  men- 
tion the  emotional  element  in  discussing  their  re- 
sults, and  inasmuch  as  they  admitted  that  they 
could  not  obviate  from  their  experimental  pro- 
cedure pain,  which  they  themselves  proved  was 
effective  in  causing  glycosuria,  designating  what 
they  called  "Fesselungsdiabetes"  as  "emotional 
glycosuria"  is  not  justified. 

Emotional  Glycosuria 

The  discovery  that  during  strong  emotion  adre- 
nal secretion  is  increased,  and  the  fact  that  injec- 
tion of  adrenin  gives  rise  to  glycosuria,  suggested 
that  glycosuria  might  be  called  forth  by  emotional 
excitement,  and  then  that  even  without  the  painful 
element  of  Bohm  and  Hoffmann's  experiments, 
sugar  might  be  found  in  the  urine.  The  testing  of 
this  possibility  was  undertaken  by  A.  T.  Shohl,  W. 
S.  Wright  and  myself  in  1911. 

Our  first  procedure  was  a  repetition  of  Bohm 
and  Hoffmann's  experiments,  freed  from  the 
factor  of  pain.  The  animals  (cats)  were  bound 
to  a  comfortable  holder,  which  left  the  head 
unfastened.  This  holder  I  had  used  hundreds 
of  times  in  X-ray  studies  of  digestion,  with 
many  different  animals,  without  causing  any  signs 
of  even  so  much  as  uneasiness.    Just  as  in  obser- 


INCREASE    OF    BLOOD    SUGAR        71 

vations  on  the  movements  of  the  alimentary  canal, 
however,  so  here,  the  animals  reacted  differently 
to  the  experience  of  being  confined.  Young  males 
usually  became  quite  frantic,  and  with  eyes  wide, 
pupils  dilated,  pulse  accelerated,  hairs  of  the  tail 
more  or  less  erect,  they  struggled,  snarling  and 
growling,  to  free  themselves.  Females,  on  the 
contrary,  especially  if  elderly,  were  as  a  rule 
much  more  calm,  and  resignedly  accepted  the  novel 
situation. 

According  to  differences  in  reaction  the  animals 
were  left  in  the  holder  for  periods  varying  in 
length  from  thirty  minutes  to  five  hours.  In 
order  to  insure  prompt  urination,  considerable 
quantities  of  water  were  given  by  stomach  tube 
at  the  beginning  of  the  experiment  and  in  some 
cases  again  later.  Arrangements  were  made  for 
draining  the  urine  promptly,  when  the  animal  was 
on  the  holder  or  when  afterwards  in  a  metal  metab- 
olism cage,  into  a  glass  receiver  containing  a  few 
drops  of  chloroform  to  prevent  fermentation. 
The  diet  in  all  cases  consisted  of  customary  raw 
meat  and  milk.  In  every  instance  the  urine  was 
proved  free  from  sugar  before  the  animal  was 
excited. 

In  our  series  of  observations  twelve  cats  were 
used,  and  in  every  one  a  well-marked  glycosuria 
was  developed.  The  shortest  periods  of  confine- 
ment to  the  holder  which  were  effective  were  thirty 


72  BODILY    CHANGES 

and  forty  minutes ;  the  longest  we  employed,  five 
hours.  The  average  time  required  to  bring  about 
a  glycosuria  was  less  than  an  hour  and  a  half; 
the  average  in  seven  of  the  twelve  cases  was  less 
than  forty  minutes.  In  all  cases  no  sugar  was 
found  in  the  urine  passed  on  the  day  after  the 
excitement. 

The  promptness  with  which  the  glycosuria  de- 
veloped was  directly  related  to  the  emotional  state 
of  the  animal.  Sugar  was  found  early  in  animals 
which  early  showed  signs  of  being  frightened 
or  in  a  rage,  and  much  later  in  animals  which  took 
the  experience  more  calmly. 

As  cooling  may  result  in  increased  sugar  in  the 
blood,  and  consequent  glycosuria,  the  rectal  tem- 
perature was  observed  from  time  to  time,  and  it 
was  found  to  vary  so  slightly  that  in  these  experi- 
ments it  was  a  wholly  negligible  factor.  In  one 
cat  the  rectal  temperature  fell  to  36°  C.  while  the 
animal  was  bound  and  placed  in  a  cold  room  (about 
2°  C.)  for  fifty  minutes,  but  no  sugar  appeared  in 
the  urine. 

Further  evidence  that  the  appearance  of  sugar 
in  the  urine  may  arise  purely  from  emotional  ex- 
citement was  obtained  from  three  cats  which  gave 
negative  results  when  bound  in  the  holder  for 
varying  periods  up  to  four  hours.  It  was  note- 
worthy that  these  animals  remained  calm  and 
passive  in  their  confinement.     When,  however, 


INCREASE    OF    BLOOD    SUGAR        73 

tliey  were  placed,  separately,  in  a  small  wire  cage, 
and  were  barked  at  by  an  energetic  little  dog,  that 
jumped  at  tliem  and  made  signs  of  attack,  the  cats 
became  much  excited,  they  showed  their  teeth, 
humped  their  backs,  and  growled  defiance.  This 
sham  fight  was  permitted  to  continue  for  a  half 
hour  in  each  of  the  three  cases.  In  each  case  the 
animal,  which  after  four  hours  of  bondage  had  ex- 
hil)ited  no  glycosuria,  now  had  sugar  in  the  urine. 
Pain,  cooling,  and  bondage  were  not  factors  in 
these  experiments.  The  animal  was  either  fright- 
ened or  enraged  by  the  barking  dog,  and  that  ex- 
citement was  attended  by  glycosuria. 

The  sugar  excreted  in  the  twenty-four  hours 
which  included  the  period  of  excitement  was  de- 
termined by  the  Bertrand  method.^^  It  ranged 
from  0.024  gram  to  1.93  grams,  or  from  0.008 
gram  to  0.62  gram  per  kilo  body  weight,  for  the 
twenty-four  hours'  quantity. 

The  presence  of  sugar  in  the  urine  may  be  used 
as  an  indication  of  increased  sugar  in  the  blood, 
for  unless  injury  has  been  done  to  the  cells  of 
the  kidneys,  they  do  not  permit  sugar  to  escape 
until  the  percentage  in  the  blood  has  risen  to  a 
considerable  degree.  Thus,  though  testing  the 
urine  reveals  the  instances  of  a  high  content  of 
blood  sugar,  it  does  not  show  the  fine  variations 
that  appear  when  the  blood  itself  is  examined. 
Recently  Scott  ^^  has  concluded  a  thorough  in- 


74  BODILY    CHANGES 

vestigation  of  the  variations  of  blood  sugar  in  cats, 
and  has  found  that  merely  incidental  conditions, 
producing  even  mild  excitement,  as  indicated  by 
crying  or  otherwise,  result  in  a  noticeable  rise  in 
the  amount.  Indeed,  so  sensitive  is  the  sugar-lib- 
erating mechanism  that  all  the  early  determina- 
tions of  the  "normal"  content  of  sugar  in  blood 
which  has  been  drawn  from  an  artery  or  vein  in 
the  absence  of  anesthesia,  are  of  very  doubtful 
value.  Certainly  when  care  is  taken  to  obtain 
blood  suddenly  from  a  tranquil  animal,  the  per- 
centage (0.069,  Scott;  0.088,  Pavy)  is  much  less 
than  when  the  blood  is  drawn  without  anesthesia 
(0.15,  Bohm  and  Hoffmann),  or  after  light  nar- 
cosis (0.282,  Eona  and  Takahashi^^). 

Our  observations  on  cats  have  since  been  found 
valid  for  rabbits.  Eolly  and  Oppermann,  Jacob- 
sen,  and  Hirsch  and  Eeinbach  ^'^  have  recently 
recorded  that  the  mere  handling  of  a  rabbit  pre- 
paratory to  operating  on  it  will  increase  the  per- 
centage of  blood  sugar  (in  some  cases  from  0.10 
to  0.23  and  0.27  per  cent).  Dogs  are  said  to  be 
much  less  likely  to  be  disturbed  by  the  nature  of 
their  surroundings  than  are  rabbits  and  cats. 
Nevertheless,  pain  and  excitement  are  such  funda- 
mental experiences  in  animals  that  without  much 
doubt  the  same  mechanism  is  operative  in  all  when 
these  experiences  occur.  Probably,  just  as  the 
digestion  of  dogs  is  disturbed  by  strong  emotion, 


INCREASE    OF    BLOOD    SUGAR        75 

the  blood  sugar  likewise  is  increased,  for  sym- 
pathetic impulses  occasion  both  changes.*  Gib 
has  given  an  account  of  a  bitch  that  became  much 
agitated  when  shut  up,  and  after  such  enforced 
seclusion,  but  never  otherwise,  she  excreted  small 
quantities  of  sugar  in  the  urine.^^ 

The  results  noted  in  these  lower  animals  have 
been  confirmed  in  human  beings.  One  of  my  for- 
mer students,  W.  G.  Smillie,  found  that  four  of 
nine  medical  students,  all  normally  without  sugar 
in  their  urine,  had  glycosuria  after  a  hard  exami- 
nation, and  only  one  of  the  nine  had  glycosuria 
after  an  easier  examination.  The  tests,  which 
were  positive  with  Fehling's  solution,  Nylander's 
reagent,  and  also  with  phenyl-hydrazine,  were 
made  on  the  first  urine  passed  after  the  exam- 
ination. Furthermore,  C.  H.  Fiske  and  I  ex- 
amined the  urine  of  twenty-five  members  of 
the  Harvard  University  football  squad  immedi- 
ately after  the  final  and  most  exciting  contest 
of  the  season  of  1913,  and  found  sugar  in 
twelve  cases.  Five  of  these  positive  cases 
were  among  substitutes  not  called  upon  to  enter 
the  game.    The  only  excited  spectator  of  the  Har- 

*  Since  the  foregoing  sentences  were  written  Hirsch  and 
Reinbach  have  reported  (Zeitschrift  fiir  physiologische 
Chemie,  1914,  xci,  p.  292)  a  "psychic  hyperglycemia"  in  dogs, 
that  resulted  from  fastening  the  animals  to  a  table.  The 
blood  sugar  rose  in  one  instance  from  0.11  to  0.14  per  cent, 
and  in  another  from  0.09  to  0.16  per  cent. 


76  BODILY    CHANGES 

vard  victory  whose  urine  was  examined  also  had 
a  marked  glycosuria,  which  on  the  following  day 
had  disappeared. 

Other  tests  made  on  students  before  and  after 
important  scholastic  examinations  have  been  pub- 
lished by  Folin,  Denis  and  Smillie.^*^  Of  thirty- 
four  second-year  medical  students  tested,  one  had 
sugar  before  the  examination  as  well  as  after- 
wards. Of  the  remaining  thirty-three,  six,  or  18 
per  cent,  had  small  but  unmistakable  traces  of 
sugar  in  the  urine  passed  directly  following  the 
ordeal.  A  similar  study  was  made  on  second-year 
students  at  a  women's  college.  Of  thirty-six  stu- 
dents who  had  no  sugar  in  the  urine  on  the  day 
before,  six,  or  17  per  cent,  eliminated  sugar  with 
the  urine  passed  immediately  after  the  examina- 
tion. 

From  the  foregoing  results  it  is  reasonable  to 
conclude  that  just  as  in  the  cat,  dog,  and  rabbit, 
so  also  in  man,  emotional  excitement  produces  tem- 
porary increase  of  blood  sugar. 

The  Eole  op  the  Adrenal  Glands  in  Emotional 
Glycosuria 

Since  artificial  stimulation  of  the  splanchnic 
nerves  produces  glycosuria,^"  and  since  major 
emotions,  such  as  rage  and  fright,  are  attended  by 
nervous  discharges  along  splanchnic  pathways, 
glycosuria  as  an  accompaniment  of  emotional  ex- 


INCREASE    OF    BLOOD    SUGAR        77 

citement  would  naturally  be  expected  to  occur. 
To  what  extent  the  adrenal  glands  which,  as 
already  mentioned,  are  stimulated  to  increased 
secretion  by  excitement,  might  play  a  part  in  this 
process,  has  been  in  dispute.  Removal  of  these 
glands  or  cutting  of  the  nerve  fibres  supplying 
them,  according  to  some  observers,-^-  prevents 
glycosuria  after  puncture  of  the  fourth  ventricle 
of  the  brain  (the  "sugar  puncture,"  which  typically 
induces  glycosuria)  and  also  after  stimulation  of 
the  splanchnics,--  On  the  other  hand,  Wert- 
heimer  and  Battez  ^^  have  stated  that  removal 
of  the  glands  does  not  abolish  the  effects  of  sugar 
puncture  in  the  cat.  It  was  questionable,  there- 
fore, whether  removal  of  the  adrenal  glands  would 
affect  emotional  glycosuria. 

Evidence  on  this  point  I  secured  with  Shohl  and 
Wright  in  observations  on  three  animals  in  which 
the  adrenals  were  removed  aseptically  under  ether. 
The  animals  selected  had  all  become  quickly  ex- 
cited on  Ijeing  bound  to  the  holder,  and  had  mani- 
fested glycosuria  after  about  an  hour  of  confine- 
ment. In  the  operation,  to  avoid  discharge  of 
adrenin  by  handling,  the  adrenal  veins  were  first 
tied,  and  then  the  glands  freed  from  their  attach- 
ments and  removed  as  quickly  and  with  as  little 
manipulation  as  possible.  In  one  cat  the  entire 
operation  was  finished  in  twenty  minutes.  In  two 
of  the  cats  a  small  catheter  was  introduced  into  the 


78  BODILY   CHANGES 

urethra  through  an  incision,  so  that  the  bladder 
could  be  emptied  at  any  time. 

In  all  three  cases  urine  that  was  free  from 
sugar  was  obtained  soon  after  the  operation.  Al- 
though the  animals  deprived  of  their  adrenals 
manifested  a  general  lessening  of  muscular  tone, 
they  still  displayed  much  of  their  former  rage  or 
excitement  when  bound.  Indeed,  one  was  more  ex- 
cited after  removal  of  the  adrenals  than  before. 
That  the  animals  might  not  be  excessively  cooled 
they  were  kept  warm  with  coverings  or  an  elec- 
tric heating  pad.  Although  they  were  now  bound 
for  periods  from  two  to  three  times  as  long  as 
the  periods  required  formerly  to  cause  glycosuria, 
no  trace  of  sugar  was  found  in  the  urine  in  any 
instance.  The  evidence  thus  secured  tends,  there- 
fore, to  support  the  view  that  the  adrenal  glands 
perform  an  important  contributory  role  in  the 
glycosuria  resulting  from  splanchnic  stimula- 
tion. 

Possibly  the  emotional  element  is  in  part  ac- 
countable for  the  glycosuria  observed  after  pain- 
ful stimulation,  but  conditions  causing  pain  alone 
will  reasonably  explain  it.  As  we  have  already 
seen,  strong  stimulation  of  sensory  fibres  causes 
the  discharge  of  impulses  along  the  splanchnic 
nerves,  and  incidentally  calls  forth  an  increased 
secretion  of  the  adrenal  glands.  In  glycosuria  re- 
sulting from  painful  stimulation,  as  well  as  in  emo- 


INCREASE    OF    BLOOD    SUGAR        79 

tional  glycosuria,  the  adrenal  glands  may  be  es- 
sential factors. 

Later  the  evidence  will  be  given  that  sugar  is 
the  optimum  source  of  muscular  energy.  In  pass- 
ing, we  may  note  that  the  liberation  of  sugar  at 
a  time  when  great  muscular  exertion  is  likely  to 
be  demanded  of  the  organism  may  be  interpreted 
as  a  highly  interesting  instance  of  biological 
adaptation. 

EEFEEENCES 

^  Kleen :  On  Diabetes  Mellitus  and  Glycosuria,  Philadel- 
phia, 1900,  pp.  22,  37-39. 

2  Naunyn :  Der  Diabetes  Mellitus,  Vienna,  1898,  p.  72. 

^  Schultze :  Verhandlungen  der  Gesellschaf t  deutscher 
Naturforscher  und  Aerzte,  Cologne,  1908,  ii,  p.  358. 

*  Raimann :  Zeitschrif t  f iir  Heilkunde,  1902,  xxiii,  Ab- 
theilung  iii,  pp.  14,  19. 

^  Mita :  Monatshefte  fUr  Psychiatric  und  Neurologie,  1912, 
xxxii,  p.  159. 

®  Folin,  Denis  and  Smillie :  Journal  of  Biological  Chem- 
istry, 1914,  xvii,  p.  519. 

^  Arndt :  Zeitschrift  fiir  Nervenheilkunde,  1897,  x.  p.  436. 

*  Naunyn:  Loc.  cit.,  p.  73;  Hirschfeld:  Die  Zuckerkrank- 
heit,  Leipzig,  1902,  p.  45. 

^  Waterman  and  Smit :  Archiv  fiir  die  gesammte  Physi- 
ologie,  1908,  cxxiv,  p.  205. 

'•'Henderson  and  Underhill :  American  Journal  of  Physi- 
ology, 1911,  xxviii,  p.  270. 

^^  Bohm  and  Hoffmann :  Archiv  fiir  experimentelle  Pa- 
thologie  und  Pharmakologie,  1878,  viii,  p.  295. 

"  Eckhard :  Zeitschrift  fiir  Biologic,  1903,  xliv,  p.  408. 

'^  Loe\vy  and  Rosenberg:  Eiochemische  Zeitschrift,  1913, 
Ivi,  p.  114. 

^*  See  Abderhaldcn :  Handbuch  der  biochemischen  Ar- 
beitsmethoden,  Berlin,  1910,  ii,  p.  181. 


80  BODILY    CHANGES 

^^  Scott :  American  Journal  of  Physiology,  1914,  sxxiv, 
p.  283. 

16  Cited  by  Scott :  Loc.  cit.,  p.  296. 

1''  Holly  and  Oppermann :  Biochemische  Zeitschrift,  1913, 
xlix,  p.  201.  Jacobsen:  Ihid.,  1913,  li,  p.  449.  Hirsch  and 
Eeinbach:  Zeitschrift  fiir  physiologische  Chemie,  1913, 
Ixxxvii,  p.  122. 

18  Cited  by  Kleen :  Loc.  cit.,  p.  37. 

1^  Folin,  Denis  and  Smillie :  Loc.   cit.,  p.   520. 

^°  See  Macleod :  American  Journal  of  Physiology,  1907, 
xix,  p.  405,  also  for  other  references  to  literature. 

^1  See  Meyer:  Comptes  rendus  de  la  Societe  de  Biologie, 
1906,  Iviii,  p.  1123;  Nishi:  Archiv  fiir  experimentelle  Pa- 
thologic und  Pharmakologie,  1909,  Ixi,  p.  416. 

^^  Gautrelet  and  Thomas :  Comptes  rendus  de  la  So- 
ciete de  Biologie,  1909,  Ixvii,  p.  233;  and  Macleod:  Pro- 
ceedings of  the  Society  for  Experimental  Biology  and  Medi- 
cine, 1911,  viii,  p.  110  (true  for  left  adrenal  and  left  splanch- 
nic), 

23  Wertheimer  and  Battez :  Archives  Internationales  de 
Physiologic,  1910,  ix,  p.  392. 


CHAPTER   VI 

IMPEOVED    CONTRACTION    OF    FATIGUED 

MUSCLE  AFTER  SPLANCHNIC   STIMULATION  OF 

THE   ADRENAL   GLAND 

In  the  older  literature  on  the  adrenal  glands  the 
deleterious  effect  of  their  absence,  or  the  beneficial 
effect  of  injected  extracts,  on  the  contraction  of 
skeletal  muscle  was  not  infrequently  noted.  As 
evidence  accumulated,  however,  tending  to  prove 
an  imjDortant  relation  between  the  extract  of  the 
adrenal  medulla  (adrenin)  and  the  sympathetic 
nervous  system,  the  relations  with  the  efficiency  of 
skeletal  muscle  began  to  receive  less  consideration. 

The  muscular  weakness  of  persons  suffering 
from  diseased  adrenals  (Addison's  disease)  was 
well  recognized  before  experimental  work  on  the 
glands  was  begun.  Experiments  on  ra])bits  were 
reported  in  1892  by  Albanese,^  who  showed  that 
muscles  which  were  stimulated  after  removal 
of  the  glands  were  much  more  exhausted  than 
when  stimulated  the  same  length  of  time  in  the 
same  animal  before  the  removal.    Similarly  Boi- 

81 


82  BODILY    CHANGES 

net  2  reported,  in  1895,  that  rats  recently  deprived 
of  their  adrenals  were  much  more  quickly  ex- 
hausted in  a  revolving  cage  than  were  normal 
animals. 

More  direct  evidence  of  the  favorable  influence 
of  adrenal  extract  on  skeletal  muscle  was  brought 
forward  by  Oliver  and  Schafer.^  After  inject- 
ing the  extract  subcutaneously  into  a  frog  they 
found  that  the  excised  gastrocnemius  muscle  regis- 
tered a  curve  of  contraction  about  33  per  cent 
higher  and  about  66  per  cent  longer  than  the 
corresponding  muscle  not  exposed  to  the  action 
of  the  extract.  Similar  prolongation  of  the  muscle 
curve  was  observed  after  injecting  the  extract 
intravenously  into  a  dog.  A  beneficial  effect 
of  adrenal  extract  on  fatigued  muscle,  even  when 
applied  to  the  solution  in  which  the  isolated 
muscle  was  contracting,  was  claimed  by  Dessy 
and  Grandis,^  who  studied  the  phenomenon  in  a 
salamander.*  Further  evidence  leading  to  the 
same   conclusion  was    offered  in  a  discriminat- 

*  These  earlier  investigations,  in  which  an  extract  of 
the  entire  gland  was  used,  made  no  distinction  between  the 
action  of  the  medulla  and  that  of  the  cortex.  It  may  be  that 
the  weakness  following  removal  or  disease  of  the  adrenals  is 
due  to  absence  of  the  cortex  (see  Hoskins  and  Wheelon :  Am- 
erican Journal  of  Physiology,  1914,  xxxiv,  p.  184).  Such  a 
possible  effect,  however,  should  not  be  confused  with  the 
demonstrable  influence  of  injected  adrenin  (derived  from  the 
adrenal  medulla  alone)  and  the  similar  effects  from  adrenal 
secretion  caused  by  splanchnic  stimulation. 


CONTRACTION  OF  FATIGUED  MUSCLE     83 

ing  paper  by  Panella.^  He  found  that  in  cold- 
blooded animals  the  active  principle  of  the  adre- 
nal medulla  notably  reinforced  skeletal  muscle, 
prolonging  its  ability  to  do  work,  and  improv- 
ing its  contraction  when  fatigued.  In  warm- 
blooded animals  the  same  effects  were  observed, 
but  only  after  certain  experimental  procedures, 
such  as  anesthesia  and  section  of  the  bulb,  had 
changed  them  to  a  condition  resembling  the  cold- 
blooded. 

The  foregoing  evidence  indicates  that  removal 
of  the  adrenals  has  a  debilitating  effect  on  muscu- 
lar power,  and  that  injection  of  extracts  of  the 
glands  has  an  invigorating  effect.  It  seemed  pos- 
sible, therefore,  that  increased  secretion  of  the 
adrenal  glands,  whether  from  direct  stimulation 
of  the  splanchnic  nerves  or  as  a  reflex  result  of 
pain  or  the  major  emotions,  might  act  as  a  dyna- 
mogenic  factor  in  the  performance  of  muscular 
work.  With  this  possibility  in  mind  L.  B.  Nice 
and  I  °  first  concerned  ourselves  in  a  research 
which  we  conducted  in  1912. 

The  general  plan  of  the  investigation  consisted 
primarily  in  observing  the  effect  of  stimulating 
the  splanchnic  nerves,  isolated  from  the  spina.1 
cord,  on  the  contraction  of  a  muscle  whose  nerve, 
also  isolated  from  the  spinal  cord,  was  rhyth- 
mically and  uniformly  excited  with  break  induc- 
tion shocks.    When  a  muscle  is  thus  stimulated  it 


84  BODILY   CHANGES 

at  first  responds  by  strong  contractions,  bnt  as 
time  passes  the  contractions  become  weaker,  the 
degree  of  shortening  of  the  muscle  becomes  less, 
and  in  this  state  of  lessened  efficiency  it  may  con- 
tinue for  a  long  period  to  do  work.  The  tired 
muscle  which  is  showing  continuously  and  evenly 
its  inability  to  respond  as  it  did  at  first,  is  said  to 
have  reached  the  "fatigue  level."  This  level  serves 
as  an  excellent  basis  for  testing  influences  that 
may  have  a  beneficial  etf ect  on  muscular  perform- 
ance, for  the  benefit  is  at  once  manifested  in 
greater  contraction. 

In  the  experimental  arrangement  which  we  used, 
only  a  connection  through  the  circulating  blood 
existed  between  the  splanchnic  region  and  the 
muscle — all  nervous  relations  were  severed.  Any 
change  in  muscular  ability,  therefore,  occurring 
when  the  splanchnic  nerve  is  stimulated,  must 
be  due  to  an  alteration  in  the  quantity  or  qual- 
ity of  the  blood  supplied  to  the  laboring 
muscle. 

Cats  were  used  for  most  experiments,  but  re- 
sults obtained  with  cats  were  confirmed  on  rab- 
bits and  dogs.  To  produce  anesthesia  in  the 
cats  and  rabbits,  and  at  the  same  time  to  avoid 
the  fluctuating  effects  of  ether,  urethane  (2  grams 
per  kilo  body-weight)  was  given  by  a  stomach  tube. 
The  animals  were  fastened  back  downward,  over 
an  electric  warming  pad,  to  an  animal  holder. 


CONTRACTION  OF  FATIGUED  MUSCLE     85 

Care  was  taken  to  maintain  the  body  temperature 
at  its  normal  level, throughout  each  experiment. 

The  Nerve-muscle  Preparation 

The  muscle  selected  to  be  fatigued  was  usually 
the  extensor  of  the  right  hind  foot  (the  tibialis 
anticus),  though  at  times  the  common  extensor 
muscle  of  the  digits  of  the  same  foot  was  em- 
ployed. The  anterior  tibial  nerve  which  supplies 
these  muscles  was  l)ared  for  about  two  centimeters, 
severed  toward  the  body,  and  set  in  shielded  elec- 
trodes, around  which  the  skin  was  fastened  by 
spring  clips.  Thus  the  nerve  could  be  protected, 
kept  moist,  and  stimulated  without  stimulation  of 
neighboring  structures.  By  a  small  slit  in  the  skin 
the  tendon  of  the  muscle  was  uncovered,  and  after 
a  strong  thread  was  tied  tightly  about  it,  it  was 
separated  from  its  insertion.  A  nerve-muscle 
preparation  was  thereby  made  which  was  still  con- 
nected with  its  proper  blood  supply.  The  prepa- 
ration was  fixed  firmly  to  the  animal  holder  by 
thongs  looped  around  the  hock  and  the  foot,  i.  e., 
on  either  side  of  the  slit  through  which  the  tendon 
emerged. 

The  thread  tied  to  the  tendon  was  passed  over 
a  pulley  and  down  to  a  pivoted  steel  bar  which 
bore  a  writing  point.  Both  the  pulley  and  this 
steel  writing  lever  were  sui)ported  in  a  rigid  tri- 
jjod.    In  the  earliest  experiments  the  contracting 


86  BODILY   CHANGES 

muscle  was  made  to  lift  weights  (125  to  175 
grams) ;  in  all  the  later  observations,  however, 
the  muscle  pulled  against  a  spring  attached  below 
the  steel  bar.  The  tension  of  the  spring  as  the 
muscle  began  to  lift  the  lever  away  from  the  sup- 
port was,  in  most  of  the  experiments,  110  grams, 
with  an  increase  of  10  grams  as  the  writing  point 
was  raised  4,5  millimeters.  The  magnification  of 
the  lever  was  3.8. 

The  stimuli  delivered  to  the  anterior  tibial  nerve 
were,  in  most  experiments,  single  break  shocks  of 
a  value  barely  maximal  when  applied  to  the  fresh 
preparation.  The  rate  of  stimulation  varied  be- 
IWeen  60  and  300  per  minute,  but  was  uniform 
ffi^miy  single  observation.  A  rate  which  was  found 
^iMrklly  serviceable  was  180  per  minute, 
^^^infee:^  the  anterior  tibial  nerve  contains  fibres 
§llS%^fife^^  blood-vessels,  as  well  as  fibres  causing 
(riSftbt^^fi^ti  of  skeletal  muscle,  the  possibility  had 
tWIBMc9^3idered  that  stimuli  applied  to  it  might 
^^t^QW^4^e^%?^od  supply  of  the  region.  Constric- 
i-ft)i  ^?^8^'^1^6'd  vessels  would  be  likely  to  pro- 
Sftfe&^te^^^Difef^^efrious  disturbance,  by  lessening 
the  blood  flow  to  the  muscle.  The  observations 
W^(k^^mt  %^i  Wktren,'  that  vasodilator  rather 
im^  ^^§M%^^§^W'^^hf£ects  are  produced  by 
%i^|l^£iaida^iJi«ai  ^0(§i§^epeated  at  intervals  of 
li^  fefSf^  #i§6  ^§^^&"§eQond,  reassured  us  as  to 
liK^J^P?  ^^(BteMMfii^the  blood  supply,  for 


CONTRACTION  OF  FATIGUED  MUSCLE     87 

the  rate  of  stimulation  in  our  experiments  never 
exceeded  five  per  second  and  was  usually  two  or 
three.  Furthermore,  in  using  these  different  rates 
we  have  never  noted  any  result  which  could  rea- 
sonably be  attributed  to  a  diminished  circulation. 

The  Splanchnic  Preparation 
The  splanchnic  nerves  were  stimulated  in  vari- 
ous ways.  At  first  only  the  left  splanchnics  in 
the  abdomen  were  prepared.  The  nerves,  sepa- 
rated from  the  spinal  cord,  were  placed  upon 
shielded  electrodes.  The  form  of  electrodes  which 
was  found  most  satisfactory  was  that  illustrated 


Figure  10. — The  shielded  electrodes  used  in  stimulating  the 
splanchnic  nerves.     For  description  see  text. 

in  Fig.  10.  The  instrument  was  made  of  a  round 
rod  of  hard  wood,  bevelled  to  a  point  at  one  end, 
and  grooved  on  the  two  sides.  Into  the  grooves 
were  pressed  insulated  wires  ending  in  platinum 
hooks,  which  projected  beyond  the  bevelled  sur- 
face. Around  the  rod  was  placed  an  insulating 
rubber  tube  which  was  cut  out  so  as  to  leave  the 
hooks  uncovered  when  the  tube  was  slipped  down- 
ward. 

In  applying  the  electrodes  the  left  splanchnic 
nerves  were  first  freed  from  their  surroundings 
and  tightly  ligatured  as  close  as  possible  to  their 


88  BODILY   CHANGES 

origin.  By  means  of  strong  compression  the  con- 
ductivity of  the  nerves  was  destroyed  central 
to  the  ligature.  The  electrodes  were  now  fixed 
in  place  by  thrusting  the  sharp  end  of  the  wooden 
rod  into  the  muscles  of  the  back.  This  was  so 
done  as  to  bring  the  platinum  hooks  a  few  milli- 
meters above  the  nerves.  With  a  small  seeker 
the  nerves  were  next  gently  lifted  over  the  hooks, 
and  then  the  rubber  tube  was  slipped  downward 
until  it  came  in  contact  with  the  body  wall.  Ab- 
sorbent cotton  was  packed  about  the  lower  end 
of  the  electrodes,  to  take  up  any  fluid  that  might 
appear;  and  finally  the  belly  wall  was  closed  with 
spring  clips.  The  rubber  tube  served  to  keep  the 
platinum  hooks  from  contact  with  the  muscles  of 
the  back  and  the  movable  viscera,  while  still  per- 
mitting access  to  the  nerves  which  were  to  be 
stimulated.  This  stimulating  apparatus  could  be 
quickly  applied,  and,  once  in  place,  needed  no 
further  attention.  In  some  of  the  experiments 
both  splanchnic  nerves  were  stimulated  in 
the  thorax.  The  rubber-covered  electrode  proved 
quite  as  serviceable  there  as  in  the  abdo- 
men. 

The  current  delivered  to  the  splanchnic  nerves 
was  a  rapidly  interrupted  induced  current  of  such 
strength  that  no  effects  of  spreading  were  notice- 
able. That  splanchnic  stimulation  causes  secre- 
tion of  the  adrenal  glands  has  been  proved  in 


CONTBACTION  OF  FATIGUED  MUSCLE     89 

many  different  ways  whicli  have  already  been  de- 
scribed (see  p.  41). 

The  Effects  of  Splanchnic  Stimulation  on  the 
Contraction  of  Fatigued  Muscle 

When  skeletal  muscle  is  repeatedly  stimulated 
by  a  long  series  of  rapidly  recurring  electric 
shocks,  its  strong  contractions  gradually  grow 
weaker  until  a  fairly  constant  condition  is  reached. 
The  record  then  has  an  even  top — the  muscle  has 
reached  the  "fatigue  level."  The  effect  of  splanch- 
nic stimulation  was  tried  when  the  muscle  had 
been  fatigued  to  this  stage.  The  effect  which  was 
often  obtained  by  stimulating  the  left  splanchnic 
nerves  is  shown  in  Fig.  11.  In  this  instance  the 
muscle  while  relaxed  supported  no  weight,  and 


Figure  11. — Upper  record,  contraction  of  the  tibialis 
anlicus,  80  times  a  minute,  lifting  a  weight  of  125  grams. 
Lower  record,  stimulation  of  the  left  splanchnic  nerves, 
two  minutes.     Time,  half  minutes. 

while  contracting  lifted  a  weight  of  125  grams. 
The  rate  of  stimulation  was  80  per  minute. 


(9^ 


BODILY    CHANGES 


Tlie  muscle  record  shows  a  brief  initial  rise 
from  the  fatigue  level,  followed  by  a  drop,  and 
that  in  turn  by  another,  prolonged  rise.  The  maxi- 
mum height  of  the  record  is  13.5  millimeters,  au 
increase  of  6  millimeters  over  the  height  recorded 
before  splanchnic  stimulation.  Thus  the  muscle 
was  performing  for  a  short  period  80  per  cent 
more  work  than  before  splanchnic  stimulation,  and 
for  a  considerably  longer  period  exhibited  an  in- 
termediate betterment  of  its  efficiency. 

The  First  Eise  in  the  Muscle  Record 

The  brief  first  elevation  in  the  muscle  record 
when  registered  simultaneously  with  arterial  blood 
pressure  is  observed  to  occur  at  the  same  time 


Figure  12. — Top  record,  arterial  blood 
pressure  with  membrane  manometer.  Mid- 
dle record,  contractions  of  tibialis  anticus 
loaded  with  125  grams  and  stimulated  80 
times  a  minute.  Bottom  record,  splanchnic 
stimulation  (two  minutes).  Time,  half  min- 
utes. 


CONTRACTION  OF  FATIGUED  MUSCLE     91 

with  the  sharp  initial  rise  in  the  blood-pressure 
curve  (see  Fig.  12).  The  first  sharp  rise  in  blood 
pressure  is  due  to  contraction  of  the  vessels  in 
the  area  of  distribution  of  the  splanchnic  nerves, 
for  it  does  not  appear  if  the  alimentary  canal  is 
removed,  or  if  the  celiac  axis  and  the  superior 
and  inferior  mesenteric  arteries  are  ligated.  The 
betterment  of  the  muscular  contraction  is  prob- 
ably due  directly  to  the  better  blood  supply  result- 
ing from  the  increased  pressure,  for  if  the  adrenal 
veins  are  clipped  and  the  splanchnic  nerves  are 
stimulated,  the  blood  pressure  rises  as  before  and 
at  the  same  time  there  may  be  registered  a  higher 
contraction  of  the  muscle. 

The  Prolonged  Rise  in  the  Muscle  Record 

As  Fig.  12  shows,  the  initial  quick  uplift  in  the 
blood-pressure  record  is  quickly  checked  by  a  drop. 
This  rapid  drop  does  not  appear  when  the  adrenal 
veins  are  ol)structed.  A  similar  difference  in 
blood-pressure  records  has  been  noted  before  and 
after  excision  of  the  adrenal  glands.  As  Elli- 
ott,® and  as  Lyman  and  I  ^  have  shown,  this 
sharp  drop  after  the  first  rise,  and  also  the  subse- 
quent elevation  of  blood  pressure,  are  the  conse- 
quences of  liberation  of  adrenal  secretion  into  the 
circulation.  Fig.  12  demonstrates  that  the  pro- 
longed rise  of  the  muscle  record  begins  soon  after 
this  characteristic  drop  in  blood  pressure. 


92  BODILY    CHANGES 

If  after  clips  have  been  placed  on  the  adre- 
nal veins  so  that  no  blood  passes  from  them,  the 
splanchnic  nerves  are  stimulated,  and  later  the 
clips  are  removed,  a  slight  bnt  distinct  improve- 
ment in  the  muscular  contraction  occurs.  As  in 
the  experiments  of  Young  and  Lehmann,^*^  in 
which  the  adrenal  veins  were  tied  for  a  time  and 
then  released,  the  release  of  the  blood  which  had 
been  pent  in  these  veins  was  quickly  followed  by 
a  rise  of  blood  pressure.  The  volume  of  blood 
thus  restored  to  circulation  was  too  slight  to  ac- 
count for  the  rise  of  pressure.  In  conjunction 
with  the  evidence  that  splanchnic  stimulation  calls 
forth  adrenal  secretion,  the  rise  may  reasonably  be 
attributed  to  that  secretion.  The  fact  should  be 
noted,  however,  that  in  this  instance  the  prolonged 
improvement  in  muscular  contraction  did  not  ap- 
pear until  the  adrenal  secretion  had  been  admitted 
to  the  general  circulation. 

Many  variations  in  the  improvement  of  activity 
in  fatigued  muscle  after  splanchnic  stimulation 
were  noted  in  the  course  of  our  investigation.  The 
improvement  varied  in  degree,  as  indicated  by  in- 
creased height  of  the  record.  In  some  instances 
the  height  of  contraction  was  doubled — a  better- 
ment by  100  per  cent ;  in  other  instances  the  con- 
traction after  splanchnic  stimulation  was  only  a 
small  fraction  higher  than  that  preceding  the  stim- 
ulation; and  in  still  other  instances  there  was  no 


CONTRACTION  OF  FATIGUED  MUSCLE    93 

betterment  whatever.  Never,  in  our  experience, 
were  the  augmented  contractions  equal  to  the 
original  strong  contractions  of  the  fresh  muscle. 
The  improvement  also  varied  in  degree  as  in- 
dicated by  persistence  of  effect.  In  some  in- 
stances the  muscle  returned  to  its  former  working 
level  within  four  or  five  minutes  after  splanchnic 
stimulation  ceased  (see  Fig.  11) ;  and  in  other  cases 
the  muscle  continued  working  with  greater  effi- 
ciency for  fifteen  or  twenty  minutes  after  the  stim- 
ulation. 

The  Two  Factors:    Arterial  Pressure  and  Adrenal 
Secretion 

The  evidence  just  presented  has  shown  that 
splanchnic  stimulation  improves  the  contraction  of 
fatigued  muscle.  Splanchnic  stimulation,  however, 
has  two  effects— it  increases  general  arterial  pres- 
sure and  it  also  causes  a  discharge  of  adrenin  from 
the  adrenal  glands.  The  questions  now  arise — 
Does  splanchnic  stimulation  produce  the  improve- 
ment in  muscular  contraction  by  increasing  the 
arterial  blood  pressure  and  thereby  flushing  the 
laboring  muscles  with  fresh  blood?  Or  does  the 
adrenin  liberated  by  splanchnic  stimulation  act 
itself,  specifically,  to  improve  the  muscular  con- 
traction? Or  may  the  two  factors  cooperate? 
These  questions  will  be  dealt  with  in  the  next  two 
chapters. 


94  BODILY    CHANGES 

EEFEEENCES 

^  Albanese :  Archives  Italiennes  de  Biologie,  1892,  xvii, 
p.  243. 

^  Boinet :  Comptes  rendus,  Societe  de  Biologie,  1895,  xlvii, 
pp.  273,  498. 

^  Oliver  and  Schafer :  Journal  of  Physiology,  1895,  xviii, 
p.  263.  See  also  Eadwanska,  Anzeiger  der  Akademie,  Krakau, 
1910,  pp.  728-736.  Reviewed  in  Zentralblatt  fiir  Biochemie 
•und  Biophysik,  1911,  xi,  p.  467. 

^  Dessy  and  Grandis :  Archives  Italiennes  de  Biologie, 
1904,  xli,  p.  231. 

^  Panella :  Archives  Italiennes  de  Biologie,  1907,  xlviii,  p. 
462. 

^  Cannon  and  Nice :  American  Journal  of  Physiology, 
1913,  xxxii,  p.  44. 

'^  Bowditch  and  Warren :  Journal  of  Physiology,  1886,  vii, 
p.  438. 

«  Elliott:  Journal  of  Physiology,  1912,  xliv,  p.  403. 

^  Cannon  and  Lyman :  American  Journal  of  Physiology, 
1913,  xxxi,  p.  376. 

^^  Young  and  Lehmann :  Journal  of  Physiology,  1908, 
xxxvii,  p.  liv. 


CHAPTER  VII 

THE  EFFECTS  ON  CONTEACTION  OF  FATIGUED 

MUSCLE  OF  VAEYIXG  THE  ARTERIAL 

BLOOD  PRESSURE 

That  great  excitement  is  accompanied  by  sym- 
pathetic innervations  which  increase  the  contrac- 
tion of  the  small  arteries,  render  unusually  forc- 
ible the  heart  beat,  and  consequently  raise  arterial 
pressure,  has  already  been  pointed  out  (see  p.  26). 
Indeed,  the  counsel  to  avoid  circumstances  likely 
to  lead  to  such  excitement,  which  is  given  to  per- 
sons with  hardened  arteries  or  with  weak  hearts, 
is  based  on  the  liability  of  serious  consequences, 
either  in  the  heart  or  in  the  vessels,  that  might 
arise  from  an  emotional  increase  of  pressure  in 
these  pathological  conditions.  That  great  muscu- 
lar effort  also  is  accompanied  by  heightened  arte- 
rial pressure  is  equally  well  known,  and  is  avoided 
by  persons  likely  to  be  injured  by  it.  Both  in  ex- 
citement and  in  strong  exertion  the  blood  is  forced 
in  large  degree  from  the  capacious  vessels  of  the 
abdomen  into  other  parts  of  the  body.    In  excite- 

95 


96  BODILY    CHANGES 

ment  the  abdominal  arteries  and  veins  are  con- 
tracted by  impulses  from  the  splanchnic  nerves. 
In  violent  effort  the  diaphragm  and  the  muscles 
of  the  belly  wall  are  voluntarily  and  antagonistic- 
ally contracted  in  order  to  stiffen  the  trunk  as  a 
support  for  the  arms ;  and  the  increased  abdominal 
pressure  which  results  forces  blood  out  of  that 
region  and  does  not  permit  reaccumulation.  The 
general  arterial  pressure  in  man,  as  McCurdy  ^ 
has  shown,  may  suddenly  rise  during  extreme 
physical  effort,  from  approximately  110  millime- 
ters to  180  millimeters  of  mercury. 

The   Effect  of   Increasing   Arterial   Pressure 

What  effect  the  increase  of  arterial  pressure,  re- 
sulting from  excitement  or  physical  strain,  may 
have  on  muscular  efficiency,  has  received  only 
slight  consideration.  Nice  and  I  found  there  was 
need  of  careful  study  of  the  relations  between 
arterial  pressure  and  muscular  ability,  and,  in 
1913,  one  of  my  students,  C.  M.  Gruber,  under- 
took to  make  clearer  these  relations. 

The  methods  of  anesthesia  and  stimulation  used 
by  Gruber  were  similar  to  those  described  in 
the  last  chapter.  The  arterial  blood  pressure  was 
registered  from  the  right  carotid  or  the  femoral 
artery  by  means  of  a  mercury  manometer.  A 
time  marker  indicating  half -minute  intervals  was 
placed  at  the  atmospheric  pressure  level  of  the 


FATIGUE    AND    BLOOD   PRESSURE    97 

manometer.  And  since  the  blood-pressure  style, 
the  writing-  point  of  the  mnscle  lever,  and  the  time 
signal  were  all  set  in  a  vertical  line  on  the  surface 
of  the  recording  drum,  at  any  given  muscular  con- 
traction the  height  of  blood  pressure  was  simul- 
taneously registered. 

To  increase  general  arterial  pressure  two  meth- 
ods were  used :  the  spinal  cord  was  stimulated  in 
the  cervical  region  through  platinum  electrodes,  or 
the  left  splanchnic  nerves  were  stimulated  after 
the  left  adrenal  gland  had  been  excluded  from  the 
circulation.  This  was  done  in  order  to  avoid  any 
influence  which  adrenal  secretion  might  exert.  It 
is  assumed  in  these  experiments  that  vessels  sup- 
plying active  muscles  would  be  actively  dilated,  as 
Kaufmann  ^  has  shown,  and  would,  therefore,  in 
case  of  a  general  increase  of  blood  pressure,  de- 
liver a  larger  volume  of  blood  to  the  area  they 
supply.  The  effects  of  increased  arterial  pressure 
are  illustrated  in  Figs.  13,  14  and  15.  In  the  ex- 
periment represented  in  Fig.  13,  the  rise  of  blood 
pressure  was  produced  by  stimulation  of  the  cer- 
vical cord,  and  in  Figs.  14  and  15  by  stimulation 
of  the  left  splanchnic  nerves  after  the  left  adre- 
nal gland  had  been  tied  off. 

The  original  blood  pressure  in  Fig.  13  was  120 
millimeters  of  mercury.  This  was  increased  by 
62  millimeters,  with  a  rise  of  only  8.4  per  cent  in 
the  height  of  contraction  of  the  fatigued  muscle. 


98 


BODILY    CHANGES 


Figure  13. — In  this  and  the  following 
records,  the  upper  curve  indicates  the 
blood  pressure,  the  middle  line  muscu- 
lar contraction,  and  the  lower  line  the  time 
in  30  seconds  (also  zero  blood  pressure.) 
Between  the  arrows  the  exposed  cervical 
spinal  cord  was  stimulated. 

In  Fig.  14  tlie  original  blood  pressure  was  100 
millimeters  of  mercury.    By  increasing  this  pres- 


FATIGUE    AND   BLOOD   PRESSURE    99 

sure  32  millimeters  there  resulted  simultaneous 
betterment  of  9.8  per  cent  in  the  height  of  muscu- 
lar contraction.  In  Fig.  14  B  the  arterial  pres- 
sure was  raised  26  millimeters  and  the  height  of 


A  B  C 

Figure    14. — Stimulation   of   the   left   splanchnic   nerves    (left 
adrenal  gland  tied  off)  during  the  periods  indicated  by  the  arrows. 

contraction  increased  correspondingly  7  per  cent. 
In  Fig.  14  C  no  appreciable  betterment  can  be  seen 
although  the  blood  pressure  rose  18  millimeters. 
In  Fig.  15  the  original  blood  pressure  was  low 
— 68  millimeters  of  mercur3\  This  was  increased 
in  Fig.  15  A  by  18  millimeters   (the  same  as  in 


100 


BODILY   CHANGES 


Fig.  14  C  without  effect),  and  there  resulted  an  in- 
crease of  20  per  cent  in  the  height  of  contraction. 
In  Fig.  15  B  the  pressure  was  raised  24  millime- 


.    A  B  C 

Figure  15. — During  the  periods  indicated  in  the 
time  line  the  left  splanchnic  nerves  were  stimulated. 
The  vessels  of  the  left  adrenal  gland  were  tied  off. 

ters  with  a  corresponding  increase  of  90  per  cent 
in  the  muscular  contraction ;  and  in  Fig.  15  C  30 
millimeters  with  a  betterment  of  125  per  cent. 

Comparison  of  Figs.  13,  14  and  15  reveals  that 
the  improvement  of  contraction  of  fatigued  mus- 
cle is  much  greater  when  the  blood  pressure  is 
raised,  even  slightly,  from  a  low  level,  than  when 
it  is  raised,  perhaps  to  a  very  marked  degree, 
from  a  high  level.  In  one  of  the  experiments  per- 
formed by  Nice  and  myself  the  arterial  pressure 


FATIGUE   AND   BLOOD   PRESSURE    101 

was  increased  by  splanclinic  stimulation  from  the 
low  level  of  48  millimeters  of  mercury  to  110  milli- 
meters, and  the  height  of  the  muscular  contrac- 
tions was  increased  about  sixfold  (see  Fig.  16). 


Figure  16  —The  bottom  record  (zero  of  blood  pressure)  shows 
stimulation  of  left  splanchnics;  between  the  arrows  the  pressure  was 
kept  from  rising  by  compression  of  heart. 

Results  confirming  those  described  above  were 
obtained  by  Gruber  in  a  study  of  the  effects  of 
splanchnic  stimulation  on  the  irritability  of  mus- 
cle when  fatigued.  In  a  series  of  eleven  observa- 
tions the  average  value  of  the  barely  effective 
stimulus  (the  "threshold"  stimulus)  had  to  be  in- 
creased as  the  condition  of  fatigue  developed.    It 


102  BODILY   CHANGES 

was  increased  for  the  nerve-muscle  by  25  per  cent 
and  for  the  muscle  by  75  per  cent.  The  left 
splanchnic  nerves,  disconnected  from  the  left  adre- 
nal gland,  were  now  stimulated.  The  arterial  pres- 
sure, which  had  varied  between  90  and  100  milli- 
meters of  mercury,  was  raised  at  least  40  milli- 
meters. As  a  result  of  splanchnic  stimulation 
there  was  an  average  recovery  of  42  per  cent  in 
the  nerve-muscle  and  of  46  per  cent  in  the  muscle. 
The  increased  general  blood  pressure  was  effec- 
tive, therefore,  quite  apart  from  any  possible 
action  of  adrenal  secretion,  in  largely  restoring  to 
the  fatigued  structures  their  normal  irritability. 

The  Effect  of  Decreasing  Arterial  Pressure 

Inasmuch  as  an  increase  in  arterial  pressure 
produces  an  increase  in  the  height  of  contraction 
of  fatigued  muscle,  it  is  readily  supposable  that 
a  decrease  in  the  pressure  would  have  the  oppo- 
site effect.  Such  is  the  case  only  when  the  blood  v 
pressure  falls  below  the  region  of  90  to  100  milli-  "^ 
meters  of  mercury.  Thus  if  the  arterial  pressure 
stands  at  150  millimeters  of  mercury,  it  has  to 
fall  approximately  55  to  65  millimeters  before 
causing  a  decrease  in  the  height  of  contraction. 
Fig.  17  is  the  record  of  an  experiment  in  which 
the  blood  pressure  was  lowered  by  lessening  the 
output  of  blood  from  the  heart  by  compressing  the 
thorax.    The  record  shows  that  when  the  pressure 


FATIGUE   AND   BLOOD   PRESSURE    103 

was  lowered  from  120  to  100  millimeters  of  mer- 
cury (A),  there  was  no  appreciable  decrease  in 
the  height  of  contraction;  when  lowered  to  90 


M*    ^. 


Figure  17. — The  arrows  indicate  the  points  at  which  the  thorax 
began  to  be  compressed  in  order  to  lessen  the  output  of  blood 
from  the  heart. 


millimeters  (B),  there  resulted  a  decrease  of  2.4 
per  cent;  when  to  80  millimeters  of  mercury  (C), 
a  decrease  of  7  per  cent;  and  when  to  70  milli- 
meters (D),  a  decrease  of  17.3  per  cent.  Results 
similar  to  those  represented  in  Fig.  17  were  ob- 
tained by  pulling  on  a  string  looped  about  the 


104  BODILY   CHANGES 

aorta  just  above  its  iliac  branches,  thus  lessening 
the  flow  to  the  hind  limbs. 

The  region  of  90  to  100  millimeters  of  mercury 
may  therefore  be  regarded  as  the  critical  region 
at  which  a  falling  blood  pressure  begins  to  be  ac- 
companied by  a  concurrent  lessening  of  the  effi- 
ciency of  muscular  contraction,  when  the  muscle 
is  kept  in  continued  activity.  It  is  at  that  region 
that  the  blood  flow  is  dangerously  near  to  being 
inadequate. 

An  Explanation  of  the  Effects  of  Varying  the  Arterial 
Pressure 

How  are  these  effects  of  increasing  and  decreas- 
ing the  arterial  blood  pressure  most  reasonably 
explained?  There  is  abundant  evidence  that  fa- 
tigue products  accumulate  in  a  muscle  which  is 
doing  work,  and  also  that  these  metabolites  inter- 
fere with  efficient  contraction.  As  Ranke  ^  long 
ago  demonstrated,  if  a  muscle,  deprived  of  circu- 
lating blood,  is  fatigued  to  a  standstill,  and  then 
the  circulation  is  restored,  the  muscle  again  re- 
sponds for  a  short  time  to  stimulation,  because 
the  waste  has  been  neutralized  or  swept  away  by 
the  fresh  blood.  When  the  blood  pressure  is  at 
its  normal  height  for  warm-blooded  animals 
(about  120  millimeters  of  mercury,  see  Fig.  13), 
the  flow  appears  to  be  adequate  to  wash  out  the 
depressive  metabolites,  at  least  in  the  single  muscle 


FATIGUE    AND    BLOOD   PRESSURE    105 

used  in  these  experiments,  because  a  large  rise  of 
pressure  produces  but  little  change  in  the  fatigue 
level.  On  the  other  hand,  when  the  pressure  is 
abnormally  low,  the  flow  is  inadequate,  and  the 
waste  products  are  permitted  to  accumulate  and 
clog  the  action  of  the  muscle.  Under  such  circum- 
stances a  rise  of  pressure  has  a  very  striking  bene- 
ficial effect. 

It  is  noteworthy  that  the  best  results  of  adre- 
nin  on  fatigued  muscle  reported  by  previous  ob- 
servers were  obtained  from  studies  on  cold-blooded 
animals.  In  these  animals  the  circulation  is  main- 
tained normally  by  an  arterial  pressure  about  one- 
third  that  of  waiTQ-blooded  animals.  Injection  of 
adrenin  in  an  amount  which  would  not  shut  off  the 
blood  supply  would,  by  greatly  raising  the  arterial 
pressure,  markedly  increase  the  circulation  of 
blood  in  the  active  muscle.  In  short,  the  conditions 
in  cold-blooded  animals  are  quite  like  those  in  the 
pithed  mammal  with  an  arterial  pressure  of  about 
50  millimeters  of  mercury  (see  Fig,  16),  Under 
these  conditions  the  improved  circulation  causes 
a  remarkable  recovery  from  fatigue.  That  notable 
results  of  adrenin  on  fatigue  are  observed  in 
warm-blooded  animals  only  when  they  are  deeply 
anaesthetized  or  are  deprived  of  the  medulla  was 
claimed  by  Panella.^  He  apparently  believed 
that  in  normal  mammalian  conditions  adrenin  has 
little  effect  because  quickly  destroyed,  whereas  in 


106  BODILY   CHANGES 

the  cold-blooded  animals,  and  in  mammals  whose 
respiratory,  circulatory,  and  thermogenic  states 
are  made  similar  to  the  cold-blooded  by  anaesthesia 
or  pithing,  the  contrary  is  true.  In  accordance 
with  our  observations  of  the  effects  of  blood  pres- 
sure on  fatigued  muscle,  we  would  explain  Panel- 
la's  results  not  as  he  has  done  but  as  due  to  two 
factors.  First,  the  efficiency  of  the  muscle,  when 
blood  pressure  is  low,  follows  the  ups  and  downs 
of  pressure  much  more  directly  than  when  the 
pressure  is  high.  And  second,  a  given  dose  of 
adrenin  always  raises  a  low  blood  pressure  in 
atonic  vessels.  The  improvement  of  circulation 
is  capable  of  explaining,  therefore,  the  main  re- 
sults obtained  in  cold-blooded  animals  and  in 
pithed  mammals. 

Oliver  and  Schafer  reported  unusually  effective 
contractions  in  muscles  removed  from  the  body 
after  adrenal  extract  had  been  injected.  As  shown 
in  Fig.  16,  however,  the  fact  that  the  circulation 
had  been  improved  results  in  continued  greater  effi- 
ciency of  the  contracting  muscle.  Oliver  and  Scha- 
f er's  observation  may  reasonably  be  accounted  for ' 
on  this  basis. 

The  "Value  of  Increased  Arterial  Pressure  in  Pain  and 
Strong  Emotion 

As  stated  in  a  previous  paragraph,  there  is  evi- 
dence that  the  vessels  supplying  a  muscle  dilate 


FATIGUE    AND   BLOOD   PRESSURE    107 

when  the  muscle  becomes  active.  And  although 
the  normal  blood  j)ressure  (about  120  millimeters 
of  mercury)  may  be  able  to  keep  adequately  sup- 
plied with  blood  the  single  muscle  used  in  our  in- 
vestigation, a  higher  pressure  might  be  required 
when  more  muscles  are  involved  in  activity,  for 
a  more  widely  spread  dilation  might  then  reduce 
the  pressure  to  the  point  at  which  there  would  be 
insufficient  circulation  in  active  organs.  Further- 
more, with  many  muscles  active,  the  amount  of 
waste  would  be  greatly  augmented,  and  the  need 
for  abundant  blood  supply  would  thereby  to  a 
like  degree  be  increased.  For  both  reasons  a  rise 
of  general  arterial  pressure  would  prove  advan- 
tageous. The  high  pressure  developed  in  excite- 
ment and  pain,  therefore,  might  be  specially  ser- 
viceable in  the  muscular  activities  which  are  likely 
to  accompany  excitement  and  pain. 

In  connection  with  the  foregoing  considerations, 
the  action  of  adrenin  on  the  distribution  of  blood 
in  the  body  is  highly  interesting.  By  measuring 
alterations  in  the  volume  of  various  viscera  and 
the  limbs,  Oliver  and  Schiifer  ■'  proved  that  the 
viscera  of  the  splanchnic  area — e.  g.,  the  spleen, 
the  kidneys,  and  the  intestines — suffer  a  consider- 
able decrease  of  volume  when  adrenin  is  adminis- 
tered, whereas  the  limbs  into  which  the  blood  is 
forced  from  the  splanchnic  region  actually  in- 
crease in  size.    The  action  of  adrenin  indicates  the 


108  BODILY   CHANGES 

relative  degrees  of  sympathetic  innervations.  In 
other  words,  at  times  of  pain  and  excitement  sym- 
pathetic discharges,  probably  aided  by  the  adrenal 
secretion  simultaneously  liberated,  will  drive  the 
blood  out  of  the  vegetative  organs  of  the  interior, 
which  serve  the  routine  needs  of  the  body,  into 
the  skeletal  muscles  which  have  to  meet  by  extra 
action  the  urgent  demands  of  struggle  or  escape. 
But  there  are  exceptions  to  the  general  state- 
ment that  by  adrenin  the  viscera  are  emptied  of 
their  blood.  It  is  well  known  that  adrenin  has  a 
vasodilator,  not  a  vasoconstrictor,  action  on  the 
arteries  of  the  heart;  it  is  well  known  also  that 
adrenin  affects  the  vessels  of  the  brain  and  the 
lungs  only  slightly  if  at  all.  From  this  evidence 
we  may  infer  that  sympathetic  impulses,  though 
causing  constriction  of  the  arteries  of  the  abdomi- 
nal viscera,  have  no  effective  influence  on  those  of 
the  pulmonary  and  intracranial  areas  and  actually 
increase  the  blood  supply  to  the  heart.  Thus  the 
absolutely  and  immediately  essential  organs — 
those  the  ancients  called  the  "tripod  of  life" — the 
heart,  the  lungs,  the  brain  (as  well  as  its  instru- 
ments, the  skeletal  muscles) — are  in  times  of  ex- 
citement abundantly  supplied  with  blood  taken 
from  organs  of  less  importance  in  critical  mo- 
ments. This  shifting  of  the  blood  so  that  there  is 
an  assured  adequate  supply  to  structures  essential 
for  the  preservation  of  the  individual  may  reason- 


FATIGUE    AND    BLOOD   PRESSURE    109 

ably  be  interpreted  as  a  fact  of  prime  biological 
significance.  It  will  be  placed  in  its  proper  setting 
when  the  other  evidence  of  bodily  changes  in  pain 
and  excitement  have  been  presented. 

REFEREXCES 

^McCurdy:  American  Journal  of  Physiology,  1901,  v, 
p.  98. 

-  Kauf mann :  Archives  de  Physiologic,  1892,  xxiv,  p.  283. 

3Ranke:  Archiv  fiir  Anatomic,  1863,  p.  446. 

*  Panella :  Archives  Italiennes  de  Biologic,  1907,  xlviii, 
p.  462. 

""  Oliver  and  Schiifer :  Journal  of  Physiology,  1895,  xviii, 
p.  240. 


CHAPTER  VIII 

THE    SPECIFIC   EOLE    OE    ADRENIN   IN 
COUNTERACTING    THE    EFFECTS    OF    FATIGUE 

As  a  muscle  approaches  its  fatigue  level,  its  con- 
tractions are  decreased  in  height.  Higher  contrac- 
tions will  again  be  elicited  if  the  stimulus  is  in- 
creased. Although  these  phenomena  are  well 
known,  no  adequate  analysis  of  their  causes  has 
been  advanced.  A  number  of  factors  are  probably 
operative  in  decreasing  the  height  of  contraction: 
(1)  The  using  up  of  available  energy-producing 
material;  (2)  the  accumulation  of  metabolites  in 
the  fatigued  muscle;  (3)  polarization  of  the  nerve 
at  the  point  of  repeated  electrical  stimulation ;  and 
(4)  a  decrease  of  irritability.  It  may  be  that  there 
are  interactions  between  these  factors  within  the 
muscle,  e.  g.,  the  second  may  cause  the  fourth. 

Variations  of  the  Threshold  Stimulus  as  a  Measure  of 
Irritability 

The  last  of  the  factors  mentioned  above — the 
effect  of  fatigue  on  the  irritability  of  the  nerve- 
muscle  combination,  or  on  the  muscle  alone — can 

110 


FATIGUE    AND   ADEENIN  111 

be  tested  by  determining  variations  in  the  least 
stimulus  capable  of  causing  the  slightest  contrac- 
tion, the  so-called  "threshold  stimulus."  As  the 
irritability  lessens,  the  threshold  stimulus  must 
necessarily  be  higher.  The  height  of  the  threshold 
is  therefore  a  measure  of  irritability.  How  does 
fatigue  alfect  the  irritability  of  nerve-muscle  and 
muscle  ?  How  is  the  irritability  of  fatigued  struc- 
tures affected  by  rest?  How  is  it  influenced  by 
adrenin  or  by  adrenal  secretion?  Answers  to 
these  questions  were  sought  in  researches  carried 
on  by  C.  M.  Gruber  ^  in  1913. 

The  Method  of  Determining  the  Threshold  Stimulus 

The  neuro-muscular  arrangements  used  in  these 
researches  were  in  many  respects  similar  to  those 
already  described  in  the  account  of  experiments 
by  Nice  and  myself.  To  avoid  the  influence  of  an 
anesthetic  some  of  the  animals  were  decerebrated 
under  ether  and  then  used  as  in  the  experiments  in 
which  urethane  was  the  anesthetic.  The  nerve 
(the  peroneus  cofnmunis)  supplying  the  tibialis  an- 
ticus  muscle  was  bared  and  severed ;  and  near  the 
cut  end  shielded  jjlatinum  electrodes  were  applied. 
These  electrodes  were  used  in  fatiguing  the  muscle. 
Between  these  electrodes  and  the  muscle  other 
platinum  electrodes  could  be  quickly  applied  to  de- 
termine the  threshold  stimulus  and  the  tissue  re- 
sistance.   These  second  electrodes  were  removed 


112  BODILY   CHANGES 

except  when  in  use,  and  when  replaced  were  set 
always  in  the  same  position.  Care  was  taken,  be- 
fore replacing  them,  to  wipe  off  moisture  on  the 
nerve  or  on  the  platinum  points. 

For  determining  the  threshold  stimulus  of  the 
muscle  the  skin  and  other  overlying  tissues  were 
cut  away  from  the  tibialis  anticus  in  two  places 
about  5  centimeters  apart.  Through  these  open- 
ings platinum  needle  electrodes  could  be  thrust 
into  the  muscle  whenever  readings  were  to  be 
taken.  Local  polarization  was  avoided  by  rein- 
serting the  needles  into  fresh  jDoints  on  the  exposed 
areas  whenever  new  readings  were  to  be  taken. 

The  tendon  of  the  tibialis  anticus  was  attached, 
as  in  the  previous  experiments,  by  a  strong  thread 
passing  about  pulleys  to  a  lever  which  when  lifted 
stretched  a  spring.  During  the  determination  of 
the  threshold  the  spring  was  detached  from  the 
lever,  so  that  only  the  pull  of  the  lever  itself 
(about  15  grams)  was  exerted  on  the  muscle. 

The  method  of  measuring  the  stimulating  value 
of  the  electric  current  which  was  used  in  testing 
the  threshold  was  that  devised  by  E.  Gr.  Martin*  of 
the  Harvard  Laboratory — a  method  by  which  the 
strength  of  an  induced  electric  shock  is  calculable 
in  definite  units.    If  the  tissue  resistance  enters 

*  For  a  full  account  of  Dr.  Martin's  method  of  calculating 
the  strength  of  electric  stimuli,  see  Martin :  The  Measurement 
of  Induction  Shocks,  New  York,  1912. 


FATIGUE    AND    ADRENIN  113 

into  the  calculation  these  are  called  /3  units.  When 
the  threshold  of  the  nen^e-muscle  was  taken,  the 
apparatus  for  the  determination  was  connected 
with  the  nerve  through  the  electrodes  nearer  the 
muscle.  They  were  separated  from  the  fatiguing 
electrodes  by  more  than  3  centimeters,  and  ar- 
ranged so  that  the  kathode  was  next  the  muscle. 
When  the  threshold  of  the  muscle  was  taken  direct- 
ly the  apparatus  was  connected  with  the  muscle 
through  platinum  needle  electrodes  thrust  into  it. 
The  position  of  the  secondary  coil  of  the  inducto- 
rium,  in  every  case,  was  read  by  moving  it  away 
from  the  primary  coil  until  the  very  smallest  pos- 
sible contraction  of  the  muscle  was  obtained.  Four 
of  these  readings  were  made,  one  with  tissue  resist- 
ance, the  others  with  10,000,  20,000,  and  30,000 
ohms  additional  resistance  in  the  secondary  cir- 
cuit. Only  break  shocks  were  employed — the 
make  shocks  were  short-circuited.  Immediately 
after  the  determination  of  the  position  of  the  sec- 
ondary coil,  and  before  the  electrodes  were  re- 
moved or  disconnected,  three  readings  of  the  tis- 
sue resistance  were  made.  From  these  data  four 
values  for  /S  were  calculated. 

The  strength  of  the  primary  current  for  deter- 
mining the  threshold  of  the  nerve-muscle  was  usu- 
ally .01  ampere,  but  in  a  few  cases  .05  ampere  was 
used.  For  normal  muscle  it  was  .05  ampere  and 
for  denervated  muscle  1.0  ampere.     The  iuducto- 


114  BODILY    CHANGES 

rium,  which  was  used  throughout,  had  a  secondary 
resistance  of  1400  ohms.  This  was  added  to  the 
average  tissue  resistance  in  making  corrections — 
corrections  were  made  also  for  core  magnetiza- 
tion. 

The  Lessening  of  Neuro-musoular  Irritability  by  Fatigue 
The  threshold  for  the  peroneus  communis  nerve 
in  decerebrate  animals  varied  from  0.319  to  2.96 
units,  with  an  average  in  sixteen  experiments  of 
1.179.*  This  average  is  the  same  as  that  found  by 
E.  L.  Porter  ^  for  the  radial  nerve  in  the  spinal 
cat.  For  animals  under  urethane  anesthesia  a 
higher  average  was  obtained.  In  these  it  varied 
from  .644  to  7.05,  or  an  average  in  ten  experiments 
of  3.081. 

The  threshold  for  the  tibialis  anticus  muscle 
varied  in  the  decerebrate  animals  from  6.75  units 
to  33.07,  or  an  average  in  fifteen  experiments  of 
18.8.  Ten  experiments  were  performed  under  ure- 
thane anesthesia  and  the  threshold  varied  from 
12.53  to  54.9,  with  an  average  of  29.84  /3  units. 
From  these  results  it  is  evident  that  anesthesia 
notably  affects  the  threshold. 

E.  L.  Porter  proved,  by  experiments  carried  on 
in  the  Harvard  Physiological  Laboratory,  that  the 
threshold  of  an  undisturbed  nerve-muscle  remains 

*  For  the  detailed  data  of  these  and  other  quantitative  ex- 
periments, the  reader  should  consult  the  tables  in  the  original 
papers. 


FATIGUE    AND    ADRENIN  115 

constant  for  hours,  and  his  observation  was  con- 
firmed by  Gruber  (see  Pig.  19).  If,  therefore, 
after  fatigue,  a  change  exists  in  the  threshold,  this 
change  is  necessarily  the  result  of  alterations  set 
up  by  the  fatigue  process  in  the  nerve-muscle  or 
muscle. 

After  fatigue  the  threshold  of  the  nerve-muscle, 
in  sixteen  decerebrate  animals,  increased  from  an 
average  of  1.179  to  3.34 — an  increase  of  183  per 
cent.  In  ten  animals  under  urethane  anesthesia 
the  threshold  after  fatigue  increased  from  a  nor- 
mal average  of  3.08  to  9.408 — an  increase  of  208 
per  cent. 

An  equal  increase  in  the  threshold  stimulus  was 
obtained  from  the  normal  muscle  directly.  In  de- 
cerebrate animals  the  normal  threshold  of  18.8 
units  was  increased  by  fatigue  to  69.54,  or  an  in- 
crease of  274  per  cent.  With  urethane  anesthesia 
the  threshold  increased  from  29.849  to  66.238,  or 
an  increase  of  122  per  cent. 

Fig.  18,  plotted  from  the  data  of  one  of  the  many 
experiments,  shows  the  relative  heights  of  the 
threshold  before  and  after  fatigue.  The  corre- 
spondence of  the  two  readings  of  the  threshold,  one 
from  the  nerve  supplying  the  muscle  and  the  other 
from  the  muscle  directly,  served  as  a  check  on  the 
electrodes.  The  broken  line  in  the  figure  repre- 
sents the  threshold  (in  units)  of  the  nerve-muscle, 
and  the  continuous  line  that  of  the  muscle.     The 


116  BODILY   CHANGES 

threshold  values  of  the  nerve-muscle  have  been 
magnified  ten  times  in  order  to  bring  the  two  rec- 
ords close  together.    In  this  experiment  the  thresh- 


FiGURE  18. — A  record  plotted  from  the  data  of  one  experiment. 
The  time  intervals  in  minutes  are  registered  on  the  abscissa;  the 
value  of  the  threshold  in  units  is  registered  on  the  ordinate.  The 
continuous  line  is  the  record  of  the  muscle,  the  broken  line  that  of 
the  nerve-muscle.  The  values  for  the  nerve-muscle  have  been 
magnified  ten  times,  those  for  the  muscle  are  normal. 

(1)  Normal  values  of  the  threshold. 

(2)  Fatigue  thresholds  after  one  hour's  work,  Ufting  120  grams 
240  times  a  minute. 

(3  and  4)  The  threshold  after  rest. 

old  of  the  muscle  after  fatigue  (i.e.,  at  2)  is  167  per 
cent  higher  than  the  normal  threshold  (at  1),  while 
that  of  the  nerve-muscle  after  fatigue  is  30.5  per 
cent  higher  than  its  normal. 

Evidently  a  direct  relation  exists  between  the 
duration  of  work  and  the  increase  of  threshold. 
For  instance,  the  threshold  is  higher  after  a  muscle 
is  fatigued  for  two  hours  than  it  is  at  the  end  of 


FATIGUE    AND    ADRENIN  117 

the  first  hour.  The  relation  between  the  work 
done  and  the  threshold  is  not  so  clear.  In  some 
animals  the  thresholds  were  higher  after  120  grams 
had  been  lifted  120  times  a  minute  for  30  minutes 
than  they  were  in  others  in  which  200  grams  had 
been  lifted  240  times  a  minute  for  the  same  period. 
The  muscle  in  the  latter  instances  did  almost  four 
times  as  much  work,  yet  the  threshold  was  lower. 
The  difference  may  be  due  to  the  general  condi- 
tion of  the  animal. 

A  few  experiments  were  performed  on  animals 
in  which  the  nerve  supplying  the  muscle  was  cut 
seven  to  fourteen  days  previous  to  the  experiment. 
The  muscle,  therefore,  had  within  it  no  living 
nerve  fibres.  The  average  normal  threshold  for 
the  denervated  muscle  in  6  animals  was  61.28  units. 
As  in  the  normal  muscle,  the  percentage  increase 
due  to  fatigue  was  large. 

The  Slow  Eestoration  of  Fatigued   Muscle  to  Normal 
Irritability  by  Rest 

That  rest  decreases  the  fatigue  threshold  of  both 
nerve-muscle  and  muscle  can  be  seen  in  Fig.  18. 
The  time  taken  for  total  recovery,  however,  is  de- 
pendent upon  the  amount  of  work  done,  but  this 
change,  like  that  of  fatigue,  varies  widely  with 
different  individuals.  In  some  animals  the  thresh- 
old returned  to  normal  in  15  minutes;  in  others, 
in  which  the  same  amount  of  work  was  done,  it  was 


118  BODILY   CHANGES 

still  above  normal  even  after  2  hours  of  rest.  This 
may  be  due  to  the  condition  of  the  animals — in 
some  the  metabolites  are  probably  eliminated  more 
rapidly  than  in  others.  There  were  also  variations 
in  the  rate  of  restoration  of  the  normal  threshold 
when  tested  on  the  nerve  and  when  tested  on  the 
muscle  in  the  same  animal.  In  Fig.  18  (at  3)  the 
nerve-muscle  returned  to  normal  in  30  minutes, 
whereas  the  muscle  (at  4)  after  an  hour's  rest  had 
not  returned  to  normal  by  a  few  /3  units.  This, 
however,  is  not  typical  of  all  nerve-muscles  and 
muscles.  The  opposite  condition — that  in  which 
the  muscle  returned  to  normal  before  the  nerve- 
muscle — occurred  in  as  many  cases  as  did  the  con- 
dition just  cited.  The  failure  of  the  two  tissues  to 
alter  uniformly  in  the  same  direction  may  be  ex- 
plained as  due  to  variations  in  the  location  of  the 
electrodes  when  thrust  into  the  muscle  at  different 
times  (e.  g.,  whether  near  nerve  filaments  or  not). 
The  results  from  observations  made  on  the  nerve 
are  more  likely  to  be  uniform  and  reliable  than  are 
those  from  the  muscle. 

The  time  required  for  the  restoration  of  the 
threshold  from  fatigue  to  normal,  in  denervated 
muscles,  is  approximately  the  same  as  that  for  the 
normal  muscle. 


FATIGUE    AND    ADRENIN  119 

The  Quick  Eestoration  of  Fatigued  Muscle  to  Normal 
Irritability  ry  Adrenin 

Tfe^  foregoing  observations  showed  that  fatigue 
'  raises  the  normal  threshold  of  a  muscle,  on  the  av- 
erage, between  100  and  200  per  cent  (it  may  be  in- 
creased more  than  600  per  cent) ;  that  this  increase 
is  dependent  on  the  time  the  muscle  works,  but 
also  varies  with  the  animal ;  that  rest,  15  minutes  to 
2  hours,  restores  the  normal  irritability;  and  that 
this  recovery  of  the  threshold  depends  upon  the 
time  given  to  rest,  the  duration  of  the  work,  and 
also  upon  the  condition  of  the  animal.  The  prob- 
lem which  was  next  attacked  by  Gruber  was  that  of 
learning  whether  the  higher  contractions  of  fa- 
tigued muscle  after  splanchnic  stimulation  could 
be  attributed  to  any  influence  which  adrenal  secre- 
tion might  have  in  restoring  the  normal  irritability. 
To  gain  insight  into  the  probabilities  he  tried  first 
the  effects  of  injecting  slowly  into  the  jugular  vein 
physiological  amounts  of  adrenin.* 

The  normal  threshold  of  the  peroneus  communis 
nerve  varied  in  the  animals  used  in  this  series  of 
observations  from  0,35  to  5.45  units,  with  an  aver- 
age in  nine  experiments  of  1.3,  a  figure  close  to  the 
1.179  found  in  the  earlier  series  on  the  effect  of 
fatigue.  For  the  tibialis  anticus  muscle,  in  which 
the  nerve-endings  were  intact,  the  threshold  varied 

•  The  form  of  adrenin  used  in  these  and  in  other  injections 
was  fresh  adrenalin  made  by  Parke,  Davis  &  Co. 


120  BODILY   CHANGES 

from  6.75  to  49.3  units,  with  an  average  in  the  nine 
experiments  of  22.2.  This  is  slightly  higher  than 
that  cited  for  this  same  muscle  in  the  earlier  series. 
By  fatigue  the  threshold  of  the  nerve-muscle  was 
increased  from  an  average  of  1.3  to  an  average  of 
3.3  units,  an  increase  of  154  per  cent.  The  muscle 
increased  from  an  average  of  22.2  to  an  average  of 
59.6,  an  increase  of  169  per  cent.  After  an  injec- 
tion of  0.1  to  0.5  cubic  centimeters  of  adrenin 
(1:100,000)  the  fatigue  threshold  was  decreased 
within  five  minutes  in  the  nerve-muscle  from  an 
average  of  3.3  to  1.8,  a  recovery  of  75  per  cent,  and 
in  the  muscle  from  an  average  of  59.6  to  42.4,  a  re- 
covery of  46  per  cent.  To  prove  that  this  effect  of 
adrenin  is  a  counteraction  of  the  effects  of  fatigue, 
Gruber  determined  the  threshold  for  muscle  and 
nerve-muscle  in  non-fatigued  animals  before  and 
after  adrenin  injection.  He  found  that  in  these 
cases  no  lowering  of  threshold  occurred,  a  result  in 
marked  contrast  with  the  pronounced  and  prompt 
lowering  induced  by  this  agent  in  muscles  when 
fatigued. 

Figs.  19  and  20,  plotted  from  the  data  of  two  of 
the  experiments,  show  the  relative  heights  of  the 
threshold  before  and  after  an  injection  of  adrenin. 
The  close  correspondence  of  the  two  readings  of 
the  threshold,  one  from  the  nerve  supplying  the 
muscle,  the  other  from  the  muscle  directly,  served 
to  show  that  there  was  no  fault  in  the  electrodes. 


FATIGUE    AND   ADRENIN  121 

The  continuous  line  in  the  Figures  represents  the 
threshold  (in  units)  of  the  muscle,  the  broken  line 
that  of  the  nerve-muscle.  The  threshold  of  the 
nerve-muscle  is  magnified  100  times  in  Fig.  19  and 
10  times  in  Fig.  20.  In  Fig.  19  (at  2  and  4)  the 
threshold  was  taken  after  an  intravenous  injection 
of  0.1  and  0.2  cubic  centimeter  of  adrenin  respec- 
tively. 

These  examples  show  that  adrenin  does  not  af- 
fect the  threshold  of  the  normal  non-fatigued  mus- 
cle when  tested  either  on  the  muscle  directly  or  on 
the  nerve-muscle.  In  Fig.  19  (at  3)  the  observa- 
tion taken  after  two  hours  of  rest  illustrates  the 
constancy  of  the  threshold  under  these  circum- 
stances. 

In  Fig.  19  the  normal  threshold  was  increased  by 
fatigue  (at  5) — the  muscle  had  been  pulling  120 
times  a  minute  for  one  hour  on  a  spring  hav- 
ing an  initial  tension  of  120  grams — from  30.0 
to  51.6  units,  an  increase  of  72  per  cent;  and  in 
the  nerve-muscle  from  0.62  to  0.89  units,  an 
increase  of  46  per  cent.  The  threshold  (at  6) 
was  taken  five  minutes  after  injecting  0.1  cubic 
centimeter  of  adrenin  (1:100,000).  The  thresh- 
old of  the  muscle  was  lowered  from  51.6  to 
38.0  units,  a  recovery  of  62  per  cent;  that  of  the 
nerve-muscle  from  0.89  to  0.79  units,  a  recovery  of 
37  per  cent.  After  another  injection  of  0.5  cubic 
centimeter  of  adrenin  the  thresholds  (at  7)  were 


122 


BODILY   CHANGES 


taken ;  that  of  the  nerve-muscle  dropped  to  normal 
— 0.59  units — a  recovery  of  100  per  cent,  and  that 


90 

- 

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I 

f        » 

— ' 

y          » 

80 

^ 

, 

/      6*^ 

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— 

^ 

\ 

70 

— 

* 

% 

- 

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~ 

...  — --orf' 

1 

601 

^"*' 

34 

i? 

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FiGtJEE  19. — A  record  plotted  from  the 
data  of  one  experiment.  The  time  mter- 
vals  m  hours  and  minutes  are  represented 
on  the  abscissa;  the  values  of  the  threshold 
in  yS  units  are  represented  on  the  ordinate. 
The  continuous  hne  is  the  record  of  the 
muscle,  the  broken  hne  that  of  the  nerv'e- 
muscle.  The  nen^e-muscle  record  is  mag- 
nified 100  times;  that  of  the  muscle  is  nor- 
mal. 

(1)  Normal  threshold  stimulus.  (2) 
Threshold  five  miautes  after  an  intraven- 
ous injection  of  0.1  cubic  centimeter  of  ad- 
renin  (1 :100,000)  without  pre^aous  fatigue. 

(3)  Threshold  after  a  rest  of  two  hours. 

(4)  Threshold  five  minutes  after  an  injec- 
tion of  0.2  cubic  centimeter  of  adrenin 
(1:100,000)  without  pre"\dous  fatigue.  (5) 
Threshold  after  one  hoiir's  fatigue.  The 
muscle  contracted  120  times  per  minute 
against  a  spring  ha\dng  an  initial  tension 
of  120  grams.  (6)  Threshold  five  minutes 
after  an  injection  (0.1  cubic  centimeter)  of 
adrenin  (1:100,000).  (7)  Threshold  five 
minutes  after  another  injection  of  adrenin 
(0.5  cubic  centimeter  of  a  1:100,000  solu- 
tion). 

of  the  muscle  remained  unaltered — 26  iDer  cent 
above  its  normal  threshold. 
Li  Fig.  20  the  threshold  (at  5)  was  taken  five 


FATIGUE    AND    ADRENIN  123 

mimites  after  an  injection  of  0.1  cubic  centimeter 
of  adrenin.  The  drop  here  was  as  large  as  that 
shown  in  Fi2-.  19.    The  threshold  taken  from  the 


- 

40 

^ 

2 

^ 

SO 

5" 

/ 

/ 

1 

to 

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2. 

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I      ^ 

7 

4 
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— e 

7 

-o 

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S:30 

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4:80 

Figure  20. — A  record  plotted  from  the  data  of  one  experiment. 
The  time  intervals  in  hours  and  minutes  are  registered  on  the 
abscissa;  the  values  of  the  threshold  in  units  are  registered  on  the 
ordinate.  The  continuous  line  is  the  record  of  the  muscle,  the 
broken  line  that  of  the  nerve-muscle.  The  record  of  the  nerve- 
muscle  is  magnified  ten  times;  that  of  the  muscle  is  normal. 

(1)  Normal  threshold.  (2)  The  threshold  after  one  hour's 
fatigue.  The  muscle  contracted  120  times  per  minute  against  a 
spring  having  an  initial  tension  of  120  grams.  (3  and  4)  Thresh- 
olds after  rest;  after  60  minutes  (3),  and  after  90  minutes  (4). 
(5)  Threshold  five  minutes  after  an  injection  of  adrenin  (0.1 
cubic  centimeter  of  a  1 :100,000  solution).  (6  and  7)  Thresholds 
after  rest;  after  60  minutes  (6),  and  after  90  minutes  (7). 

muscle  directly  was  lowered  from  30.6  to  18  units, 
a  recovery  of  61  per  cent;  the  nerve-muscle  from 
1.08  to  0.87  units,  a  recovery  of  51  per  cent.  That 
this  sudden  decrease  cannot  be  due»to  rest  is  shown 
in  the  same  Figure  (at  3  and  4).  These  readings 
were  made  after  60  and  90  minutes'  rest  respective- 
ly. The  sharp  decline  in  the  record  (at  5)  indi- 
cates distinctly  the  remarkable  restorative  influ- 


124  BODILY   CHANGES 

ence  of  adrenin  in  promptly  lowering  the  high 
fatigue  threshold  of  nenro-muscular  irritability. 

The  Evidence  that  the  Eestorative  Action  of  Adrenin  is 

Specific 

As  stated  in  describing  the  effects  of  arterial 
blood  pressure,  an  increase  of  pressure  is  capable 
of  causing  a  decided  lowering  of  the  neuro-muscu- 
lar  threshold  after  fatigue.  Is  it  not  possible  that 
adrenin  produces  its  beneficial  effects  by  better- 
ing the  circulation  ? 

Nice  and  I  had  argued  that  the  higher  contrac- 
tions of  fatigued  muscle,  that  follow  stimulation  or 
injection  of  adrenin,  could  not  be  wholly  due  to 
improved  blood  flow  through  the  muscle,  for  when 
by  traction  on  the  aorta  or  compression  of  the 
thorax  arterial  pressure  in  the  hind  legs  was  pre- 
vented from  rising,  splanchnic  stimulation  still 
caused  a  distinct  improvement,  the  initial  appear- 
ance of  which  coincided  with  the  point  in  the  blood- 
pressure  curve  at  which  evidence  of  adrenal  secre- 
tion appeared.  And,  furthermore,  the  improve- 
ment was  seen  also  when  adrenin  was  given  intra- 
venously in  such  weak  solution  (1:100,000)  as  to 
produce  a  fall  instead  of  a  rise  of  arterial  pressure. 
Lyman  and  I  had  shown  that  this  fall  of  pressure 
was  due  to  a  dilator  effect  of  adrenin.  Since  the 
blood  vessels  of  the  fatigued  muscle  were  dilated  by 
severance  of  their  nerves  when  the  nerve  trunk  was 


FATIGUE    AND    ADRENIN 


135 


cut,  and,  besides,  as  previously  stated  (see  p.  86), 
were  being  stimulated  tbrough  tbeir  nerves  at  a 
rate  favorable  to  relaxation,  it  seemed  hardly  prob- 


FiGtjRE  21. — Top  record,  blood  pressure 
with  mercury  manometer.  Middle  record, 
contractions  of  the  tibialis  anticus  muscle 
240  times  per  minute  against  a  spring  with 
an  initial  tension  of  120  grams.  Bottom 
record  (zero  blood  pressure),  injection  of 
0.4  cubic  centimeter  of  adrenin  (1:100,- 
000).     Time  in  half  minutes. 

able  that  adrenin  could  produce  its  beneficial  effect 
by  further  dilation  of  the  vessels  and  by  consequent 
flushing  of  the  muscle  with  an  extra  supply  of 
blood.2     The    lowering    of    blood    pressure    had 


126  BODILY    CHANGES 

been  proved  to  have  no  other  effect  than  to  impair 
the  action  of  the  muscle  (see  p.  103) .  Although  the 
chances  were  thus  against  an  interpretation  of  the 
beneficial  influence  of  adrenin  through  action  on 
the  circulation,  it  was  thought  desirable  to  test 
the  possibility  by  comparing  its  effect  with  that 
of  another  vasodilator — amyl  nitrite. 

Figs.  21  and  22  are  curves  obtained  from  the  left 
tibialis  anticus  muscle.  The  rate  of  stimulation 
was  240  times  a  minute. 

The  muscle  in  Fig.  21  contracted  against  a  spring 
having  an  initial  tension  of  120  grams,  and  that  in 
Fig.  22  against  an  initial  tension  of  100  grams.  In 
Fig.  21,  at  the  point  indicated  on  the  base  line,  0.4 
cubic  centimeter  of  adrenin  (1:100,000)  was  in- 
jected into  the  left  external  jugular  vein.  There 
resulted  a  fall  of  25  millimeters  of  mercury  in  the 
arterial  pressure  and  a  concurrent  betterment  of  15 
per  cent  in  the  height  of  contraction,  requiring 
two  minutes  and  fifteen  seconds  of  fatigue  (about 
540  contractions)  before  it  returned  to  the  former 
level.  In  Fig.  22,  at  the  point  indicated  by  the 
arrow,  a  solution  of  amyl  nitrite  was  injected  into 
the  right  external  jugular  vein.  There  resulted  a 
fall  of  70  millimeters  of  mercury  in  arterial  pres- 
sure and  a  betterment  of  4.1  per  cent  in  the  height 
of  muscular  contraction,  requiring  fifteen  seconds 
of  fatigue  (about  60  contractions)  to  decrease  the 
height   of   contraction   to   its   former  level.      In 


FATIGUE   AND   ADEENIN  1*29 


Figure  22. — Top  record, 
blood  pressure  with  mercury 
manometer.  Middle  record,  con- 
tractions of  tibialis  anlicus  mus- 
cle 240  per  minute  against  a 
spring  with  an  initial  tension  of 
100  grams  direct  load.  Bottom 
record  (zero  blood  pressure),  time 
in  half  minutes.  The  arrow  indi- 
cates the  point  at  which  a  solu- 
tion of  amyl  nitrite  was  injected. 


12«^  BODILY    CHANGES 

neither  case  did  the  blood  pressure  fall  below  the 
critical  region  (see  p.  104).* 

Although  the  fall  in  arterial  pressure  caused  by 
dilation  of  the  vessels  due  to  amyl  nitrite  was  al- 
most three  times  as  great  as  that  produced  by  the 
adrenin,  yet  the  resultant  betterment  was  only 
about  one-fourth  the  percentage  height  and  lasted 
but  one-ninth  the  time.  In  all  cases  in  which  these 
solutions  caused  an  equal  fall  in  arterial  pressure, 
adrenin  caused  higher  contractions,  whereas  amyl 
nitrite  caused  no  appreciable  change. 

The  Point  of  Action  of  Adrenin  in  Muscle 

From  the  evidence  presented  in  the  foregoing 
pages  it  is  clear  that  adrenin  somehow  is  able  to 
bring  about  a  rapid  recovery  of  normal  irritability 
of  muscle  after  the  irritability  has  been  much  less- 
ened by  fatigue,  and  that  the  higher  contractions 
of  a  fatigued  muscle  after  an  injection  of  adrenin 
are  due,  certainly  in  part,  to  some  specific  action 
of  this  substance  and  not  wholly  to  its  influence  on 
the  circulation.    Some  of  the  earlier  investigators 

*  In  some  cases  after  injection  of  amyl  nitrite  the  normal 
blood  pressure,  which  was  high,  dropped  sharply  to  a  point 
below  the  critical  region.  There  resulted  a  primary  increase 
in  muscular  contraction  due  to  the  betterment  in  circulation 
caused  by  the  dilation  of  the  vessels  before  the  critical  region 
was  reached.  During  the  time  that  the  pressure  was  below 
the  critical  region  the  muscle  contraction  fell.  As  the  blood 
pressure  again  rose  to  normal  the  muscle  contraction  in- 
creased coincidently. 


FATIGUE  AND  ADRENIN  129 

of  adrenal  function,  notably  Albanese,^  and  also 
Abelous  and  Langlois,^  inferred  from  experi- 
ments on  the  removal  of  the  glands  that  the  role 
they  played  in  the  bodily  economy  was  that  of  neu- 
tralizing, destroying  or  transforming  toxic  sub- 
stances" produced  in  the  organism  as  a  result  of 
muscular  or  nervous  work.  It  seemed  possible  that 
the  metabolites  might  have  a  checking  or  blocking 
influence  at  the  junction  of  the  nerve  fibres  with  the 
muscle  fibres,  and  might  thus,  like  curare,  lessen  the 
efficiency  of  the  nerve  impulses.  Radwjanska's  ob- 
servation ^  that  the  beneficial  action  of  adrenin  is 
far  greater  when  the  muscle  is  stimulated  through 
its  nerve  than  when  stimulated  directly,  and  Panel- 
la's  discovery '  that  adrenin  antagonizes  the  ef- 
fect of  curare,  were  favorable  to  the  view  that 
adrenin  improves  the  contraction  of  fatigued  mus- 
cle by  lessening  or  removing  a  block  established  by 
accumulated  metabolites. 

The  high  threshold  of  fatigued  denervated  mus- 
cle, however,  Gruber  found  was  quite  as  promptly 
lowered  by  adrenin  as  was  that  of  normal  muscles 
stimulated  through  their  nerves.  Fig,  23  shows 
that  the  height  of  contraction,  also,  of  the  fatigued 
muscle  is  increased  when  adrenin  is  administered. 
In  this  experiment  the  left  tibialis  anticus  muscle 
was  stimulated  directly  by  thrusting  platinum 
needle  electrodes  into  it.  The  pcroneus  communis 
nerve  supplying  the  muscle  had  been  cut  and  two 


130  BODILY   CHANGES 

centimeters  of  it  removed  nine  days  previous  to 
the  experiment.  The  rate  of  stimulation  was  120 
times  per  minute  and  the  initial  tension  of  the 
spring  about  120  grams.    At  the  point  indicated 


Figure  23. — Top  record,  blood  pressure  with 
mercury  manometer.  Middle  record,  contractions 
of  a  denervated  muscle  {tibialis  anticus)  240  per 
per  minute  against  a  spring  having  an  initial  ten- 
sion of  120  grams  (peroneus  communis  nerve  was 
cut  nine  days  before  this  record  was  taken).  Bot- 
tom record  (zero  blood  pressure),  time  in  half  min- 
utes. At  the  point  indicated  by  an  arrow  0.1  cubic 
centimeter  of  adrenin  (1:100,000)  was  injected 
intravenously. 

by  the  arrow  an  injection  of  0.1  cubic  centimeter 
of  adrenin  (1:100,000)  was  made  into  a  jugular 
vein.  A  fall  in  arterial  pressure  from  110  to  86 
millimeters  of  mercury  and  a  simultaneous  better- 
ment of  20  per  cent  in  the  height  of  contraction 


FATIGUE    AND   ADRENIN  131 

were  obtained.  It  required  four  minutes  of  fatigue 
(about  480  contractions)  to  restore  the  muscle 
curve  to  its  former  level.  Results  similar  to  this 
were  obtained  from  animals  in  which  the  nerve  had 
been  cut  7,  9,  12,  14,  and  21  days.  In  all  instances 
the  nerve  was  inexcitable  to  strong  faradic  stimula- 
tion. 

In  Radwanska's  experiments,  mentioned  above, 
the  muscle  was  stimulated  directly  when  the  nerve 
endings  were  intact.  It  seems  reasonable  to  sup- 
pose, therefore,  that  in  all  cases  he  was  stimulat- 
ing nerve  tissue.  Since  a  muscle  is  more  irritable 
when  stimulated  through  its  nerve  than  when 
stimulated  directly  (nerve  and  muscle),  a  slight 
change  in  the  irritability  of  the  muscle  by  adrenin 
would  naturally  result  in  a  greater  contraction 
when  the  nerve  was  stimulated.  Panella's  results 
also  are  not  inconsistent  with  the  interpretation 
that  the  effect  of  adrenin  is  on  the  muscle  substance 
rather  than  on  the  nerve  endings.  A  method  which 
has  long  been  used  to  separate  muscle  from  nerve 
is  that  of  blocking  the  nervous  impulses  by  the 
drug  curare.  Gruber  found  that  when  curare  is  in- 
jected the  threshold  of  the  normal  muscle  is  in- 
creased as  was  to  be  expected  from  the  removal  of 
the  highly  efficient  nervous  stimulations.  And  also, 
as  was  to  be  expected  on  that  basis,  curare  did  not 
increase  the  threshold  in  a  muscle  in  which  the 
nerve  endings  had  degenerated.    Adrenin  antago- 


132  '  BODILY   CHANGES 

nizes  curare  with  great  promptness,  decreasing  the 
heightened  threshold  of  a  curarized  muscle,  in  five 
minutes  or  less,  in  some  cases  to  normal.  From 
this  observation  it  might  be  supposed  that  curare 
and  fatigue  had  the  same  effect,  and  that  adrenin 
had  the  single  action  of  opposing  that  effect.  But 
fatigue  raises  the  threshold  of  a  curarized  muscle, 
and  adrenin  then  antagonizes  this  fatigue.  Lang- 
ley  ^  has  argued  that  curare  acts  upon  a  hypo- 
thetical "receptive  substance"  in  muscle.  If  so, 
probably  curare  acts  upon  a  substance,  or  at  a 
point,  different  from  that  upon  which  fatigue  acts ; 
for,  as  the  foregoing  evidence  shows,  fatigue  in- 
creases the  threshold  of  a  muscle  whether  deprived 
of  its  nerve  supply  by  nerve  section  and  degenera- 
tion or  by  curare,  whereas  curare  affects  only  the 
threshold  of  a  muscle  in  which  the  nerve  endings 
are  normal.''  And  since  adrenin  can  oppose  the 
effects  of  both  curare  and  fatigue,  it  may  be  said 
to  have  two  actions,  or  to  act  on  two  different 
substances  or  at  two  different  points  in  the  muscle. 
The  evidence  adduced  in  the  last  chapter  indi- 
cated that  the  greater  "head"  of  arterial  pressure 
produced  by  the  more  rapid  heart  beat  and  the 
stronger  contraction  of  many  arterioles  in  times  of 
great  excitement  would  be  highly  serviceable  to  the 
organism  in  any  extensive  muscular  activity  which 
the  excitement  might  involve.  By  assuring  an 
abundant  flow  of  blood  through  the  enlarged  ves- 


FATIGUE   AND   ADRENIN  133 

sels  of  the  working  muscle,  the  waste  products 
resulting  from  the  wear  and  tear  in  contraction 
would  be  promptly  swept  away  and  thus  would 
be  prevented  from  imi^airing  the  muscular  effi- 
ciency. The  adrenin  discharge  at  such  times  would, 
as  was  pointed  out,  probably  reinforce  the  effects 
of  sympathetic  impulses.  The  evidence  presented 
in  this  chapter  shows  that  adrenin  has  also  another 
action,  a  very  remarkable  action,  that  of  restor- 
ing to  a  muscle  its  original  ability  to  resj)ond  to 
stimulation,  after  that  has  been  largely  lost  by 
continued  activity  through  a  long  period.  What 
rest  will  do  only  after  an  hour  or  more,  adrenin 
will  do  in  five  minutes  or  less.  The  bearing  of  this 
striking  phenomenon  on  the  functions  of  the  or- 
ganism in  times  of  great  need  for  muscular  activ- 
ity will  be  considered  in  a  later  discussion. 

REFERENCES 

^  Gruber:  American  Journal  of  Physiology,  1913,  xxxii, 
p.  437. 

^  E.  L.  Porter :  American  Journal  of  Physiology,  1912, 
xxxi,  p.  149. 

•■'Cannon  and  Nice:  American  Journal  of  Physiology, 
1913,  xxxii,  p.  55. 

^Albanese:  Archives  Italiennes  de  Biologic,  1892,  xvii, 
p.  239. 

^  Abelous  and  Langlois :  Archives  de  Physiologic,  1892, 
xxiv,  pp.  269-278,  465-476. 

*■*  Radwanska :  Anzeiger  der  Akademie,  Krakau,  1910,  pp. 
728-736.  Reviewed  in  the  Centralblatt  fiir  Biochemie  und 
Biophysik,  1911,  xi,  p.  467. 


134  BODILY    CHANGES 

■^Panella:  Archives  Italiennes  de  Biologie,  1907,  xlvii, 
p.  30. 

*  Langley :  Proceedings  of  the  Royal  Society  of  London, 
1906,  Ixxviii,  B,  p.  181.  Journal  of  Physiology,  1905-6, 
xxxiii,  pp.  374-413. 

^  See  Gruber :  American  Journal  of  Physiology,  1914, 
xxxiv,  p.  89. 


CHAPTER  IX 

THE  HASTENING  OF   COAGULATION   OF  BLOOD 
BY   ADEENIN 

The  primary  value  of  blood  to  the  body  must 
have  been  one  of  the  earliest  observations  of  rea- 
soning beings.  When  we  consider  the  variety  of 
fundamental  services  which  this  circulating  fluid 
performs — the  conveyance  of  food  and  oxygen  to 
all  the  tissues,  the  removal  of  waste,  the  delivery  of 
the  internal  secretions,  the  protection  of  the  body 
against  toxins  and  bacterial  invasion,  and  the  dis- 
tribution of  heat  from  active  to  inactive  regions — 
the  view  of  the  ancient  Hebrews  that  the  "life  of 
the  flesh  is  in  the  blood"  is  well  justified.  It  is 
naturally  of  the  utmost  importance  that  this  pre- 
cious fluid  shall  be  safeguarded  against  loss.  And 
its  property  of  turning  to  a  jelly  soon  after  escap- 
ing from  its  natural  channels  assures  a  closure  of 
the  opening  through  which  the  escape  occurred,  and 
thus  protection  of  the  body  from  further  bleeding. 
The  slight  evidence  that  adrenin  hastens  the  clot- 
ting process  has  already  been  hinted  at.    When  we 

135 


136  BODILY   CHANGES 

found  that  adrenin  is  set  free  in  pain  and  intense 
emotion,  it  seemed  possible  that  there  might  exist 
in  the  body  an  arrangement  for  making  doubly 
sure  the  assurance  against  loss  of  blood,  a  proc- 
ess that  might  nicely  play  its  role  precisely  when 
the  greatest  need  for  it  would  be  likely  to  arise. 
It  was  in  1903,  while  tracing  in  dogs  the  rise  and 
fall  of  sugar  in  the  blood  after  administering 
adrenin,  that  Vosburgh  and  Eichards  ^  first  noted 
that  simultaneously  with  the  increase  of  blood 
sugar  there  occurred  more  rapid  coagulation.    In 
some  cases  the  diminution  was  as  much  as  four- 
fifths  the  coagulation  time  of  the  control.     Since 
this  result  was  obtained  by  painting  "adrenalin" 
on  the  pancreas,  as  well  as  by  injecting  it  into  the 
abdominal  cavity,  they  concluded  that  "the  phe- 
nomenon appears  to  be  due  to  the  application  of 
adrenalin  to  the  pancreas."    Six  years  later,  dur- 
ing a  study  of  the  effect  of  adrenalin  on  internal 
hemorrhage,  Wiggers  ^  examined  incidentally  the 
evidence  presented  by  Vosburgh  and  Eichards, 
and  after  many  tests  on  five  dogs  found  "never 
the  slightest  indication  that  adrenalin,  either  when 
injected  or  added  to  the  blood,  appreciably  hast- 
ened the  coagulation  process."     In  1911  von  den 
Velden^  reported  that  adrenin  (about  0.007  mil- 
ligram per  kilo  of  body  weight)   decreased  the 
coagulation  time  in  man  about  one-half — an  effect 
appearing   11   minutes   after   administration   by 


FASTEE  COAGULATION  BY  ADRENIN  137 

mouth,  and  85  minutes  after  subcutaneous  injec- 
tion. He  affirmed  also,  but  without  describing  the 
conditions  or  giving  figures,  that  adrenin  de- 
creases coagulation  time  in  vitro.  He  did  not  at- 
tribute the  coagulative  effect  of  adrenin  in  patients 
to  this  direct  action  on  the  blood,  however,  but  to 
vasoconstriction  disturbing  the  normal  circulation 
and  thereby  the  normal  equilibrium  between  blood 
and  tissue.  In  consequence,  the  tissue  juices  with 
their  coagulative  properties  enter  the  blood,  so  he 
assumed.  In  support  of  this  theory  he  offered  his 
observation  that  coagulation  time  is  decreased 
after  the  nasal  mucosa  has  been  rendered  anemic  by 
adrenin  pledgets.  Von  den  Velden's  claim  ^  for 
adrenin  given  by  mouth  was  subjected  to  a  single 
test  on  man  by  Dale  and  Laidlaw,^  but  their  re- 
sult was  completely  negative. 

The  importance  of  Vosburgh  and  Richards'  ob- 
servation, the  thoroughly  discordant  testimony  of 
later  investigators,  as  well  as  the  meager  and  inci- 
dental nature  of  all  the  evidence  that  has  been  ad- 
duced either  for  or  against  the  acceleration  of  clot- 
ting by  adrenin,  made  desirable  a  further  study  of 
this  matter.  Especially  was  this  further  study  de- 
sirable because  of  the  discharge  of  adrenin  into 
the  blood  in  pain  and  emotional  excitement.  Ac- 
cordingly, in  1914,  II.  Gray  and  I  ^  undertook  an 
investigation  of  the  question.  In  doing  so  we  em- 
ployed cats  as  subjects.    Usually  they  were  quickly 


138  BODILY   CHANGES  / 

decerebrated  under  ether,  and  then  continuance  of 
the  drug  became  unnecessary.  Body  temperature 
was  maintained  by  means  of  an  electric  heating 
pad.  Respiration  proceeded  normally  except  in  a 
few  instances  (in  which,  presumably,  there  was 
hemorrhage  into  the  medulla),  when  artificial  res- 
piration had  to  be  given. 

The  Graphic  Method  of  Measuring  the  Coagulation 
Time 

In  order  to  avoid,  so  far  as  possible,  the  personal 
element  in  determining  when  the  blood  was  clotted, 


a 


) 


Figure  24. — Diagram  of  the  graphic  coagulometer.  The  can- 
nula at  the  right  rests  in  a  water  bath  not  shown  in  this  diagram. 
For  further  description  see  text. 

the  blood  was  made  to  record  its  own  clotting.  The 
instrument  by  means  of  which  this  was  done  was 
the  graphic  coagulometer  devised  by  W.  L.  Men- 
denhall  and  myself,^  and  illustrated  diagram- 
matically  in  Fig.  24.  It  consists  essentially  of  a 
light  aluminum  lever  with  the  long  arm  nearly 
counterpoised  by  a  weight  W.    The  long  arm  is 


FASTER  COAGULATION  BY  ADRENIN  139 

prevented  from  falling  by  a  support  S,  and  is  pre- 
vented from  rising  by  a  horizontal  right-angled 
rod  reaching  over  the  lever  at  R^  and  fixed  into  the 
block  B  which  turns  on  the  axis  A.  Into  the  same 
block  is  fixed  the  vertical  rod  R~.  When  this  rod 
is  moved  from  the  post  P^,  against  which  it  is  held 
by  the  weight  of  the  horizontal  rod  R\  towards  the 
other  post  P-,  the  check  on  the  long  arm  of  the 
lever  is  lifted,  and  if  the  short  arm  is  heavier,  the 
long  arm  will  then  rise. 

The  cannula  C,  into  which  the  blood  is  received, 
is  two  centimeters  in  total  length  and  slightly  more 
than  two  millimeters  in  internal  diameter.  It  is 
attached  by  a  short  piece  of  rubber  tubing  to  the 
tapered  glass  tube  T,  five  centimeters  long  and  five 
millimeters  in  internal  diameter.  The  upper  end 
of  this  tube  is  surrounded  by  another  piece  of  rub- 
ber which  supports  the  tube  when  it  is  slid  into  the 
U-shaped  support  U,  fixed  directly  below  the  end 
of  the  short  arm  of  the  lever. 

By  drawing  the  cannulas  from  a  single  piece  of 
glass  tubing  and  by  making  the  distance  from 
shoulder  to  upper  end  about  twelve  millimeters, 
receptacles  of  fairly  uniform  capacity  are  assured. 
All  the  dimensions,  the  reach  of  the  rubber  con- 
nection over  the  top  of  the  cannula  (2-3  milli- 
meters), the  distance  of  the  upper  rubber  ring 
from  the  lower  end  of  the  glass  chamber  (4  centi- 
meters), etc.,  were  as  nearly  standard  as  possible. 


140  BODILY   CHANGES 

A  copper  wire  D,  eiglit  centimeters  long  and  0.6 
millimeters  in  diameter,  bent  above  into  a  book 
and  below  into  a  small  ring  sligbtly  less  tban  two 
millimeters  ia  diameter,  is  bnng  in  a  depression  at 
the  end  of  tbe  sbort  aiTQ  of  tbe  lever.  The  small 
ring  then  rests  in  tbe  npper  part  of  tbe  cannula 
(see  Fig.  24).  Tbe.  weigbt  of  tbe  copper  wire 
makes  tbe  sbort  arm  of  tbe  lever  beavier  tban  tbe 
long  arm  by  30  milligrams,  wben  tbe  delicate  writ- 
ing point  is  moving  over  a  ligbtly  smoked  drum. 
Half  a  dozen  of  tbese  standard  wires  are  needed. 

For  accurate  determination  of  tbe  coagulation 
time  Addis  '  bas  defined  tbe  following  conditions 
as  essential: 

1.  Tbe  blood  must  always  be  obtained  under  tbe 
same  conditions. 

2.  Estimates  must  all  be  made  at  tbe  same  tem- 
perature. 

3.  Tbe  blood  must  always  come  in  contact  witb 
tbe  same  amount  and  kind  of  foreign  material. 

4.  Tbe  end  point  must  be  clear  and  definite  and 
must  always  indicate  tbe  same  degree  of  coagula- 
tion. 

Tbe  precautions  taken  to  fulfill  tbese  conditions 
were  as  follows: 

1.  Drawing  the  hlood. — Tbe  blood  was  taken 
from  tbe  femoral  artery.  Tbe  artery  (usually  tbe 
rigbt)  was  laid  bare  in  tbe  groin  and  freed  from 
surrounding  tissue.     A  narrow  arteiy  clip,  witb 


FASTER  COAGULATION  BY  ADRENIX  141 

each  limb  enclosed  in  soft  rubber  tubing  (to  pre- 
vent injury  of  the  tissues),  and  with  its  spring  ex- 
erting gentle  pressure,  was  placed  on  the  artery 
immediately  below  the  deep  femoral  branch,  thus 
allowing  no  blood  to  stagnate  above  the  clip.  Be- 
tween the  clip  and  a  ligature  applied  about  1.5 
centimeters  below,  an  opening  was  made.  The 
blood  was  carefully  milked  out  of  the  vessels  be- 
tween a  blunt  dissector  moved  beneath,  and  a  small 
forceps,  twisted  into  a  pinch  of  absorbent  cotton, 
moved  above. 

The  cannula,  cleaned  in  water,  alcohol,  and  ether, 
was  set  in  the  rubber  connection  of  the  glass  tube ; 
the  point  of  the  cannula  was  then  lubricated  with 
vaseline  and  slipped  into  the  artery.  The  pres- 
sure of  the  clip  on  the  artery  was  next  very  slightly 
released  and  blood  was  allowed  to  flow  into  the 
cannula  up  to  the  lower  border  of  the  rubber  con- 
nection. Only  a  good-sized  drop  of  blood  was 
needed.  Sometimes  the  blood  ran  one  or  two  milli- 
meters above  or  below,  but  without  appreciably 
changing  the  result.  Since  the  clip  was  situated  on 
the  femoral  immediately  below  a  branch  in  which 
the  circulation  persisted,  the  blood  received  in  the 
cannula  teas  always  fresh  from  the  moving  stream. 
As  soon  as  the  clip  gripped  the  artery  again,  the 
cannula  was  slipped  out.  A  helper  then  promptly 
milked  the  vessel  in  the  manner  described  above, 
and  covered  it  with  a  pad  of  absorbent  cotton 


142  BODILY   CHANGES 

smeared  with  vaseline  to  prevent  drying.  There- 
by blood  was  not  permitted  to  stagnate;  and  when 
a  new  sample  was  to  be  taken,  the  vessel  was  clean 
and  ready  for  use. 

The  tip  of  the  cannula  was  at  once  plugged  by 
plunging  it  into  a  flat  mound  of  plasticine  about 
three  millimeters  high.  It  was  drawn  off  sidewise 
lest  the  plasticine  plug  be  pulled  out  again.  One 
of  the  copper  wires  D  was  now  slid  into  the  tube 
and  cannula,  the  tube  slipped  into  the  U-support, 
and  the  wire  lifted  and  hung  on  the  lever.  This 
procedure,  from  the  moment  blood  began  to  flow 
until  the  wire  was  hung,  consumed  usually  about 
twenty  seconds. 

2.  Uniform  temperature. — Under  the  U-support 
was  placed  a  large  water  bath,  in  which  the  can- 
nula and  the  tapering  part  of  the  tube  were  sub- 
merged. A  thermometer  was  fixed  to  the  U-sup- 
port so  that  the  bulb  came  near  the  cannula  in  the 
bath.  The  water  was  kept  within  a  degree  of  25° 
C.  This  temperature  was  chosen  for  several  rea- 
sons: (a)  The  cannula  has  room  temperature  and 
rapidly  cools  the  small  volume  of  blood  that  enters 
it.  To  heat  blood  and  cannula  to  body  tempera- 
ture would  take  time.  A  bath  near  room  tempera- 
ture, therefore,  seems  preferable  to  one  near  body 
temperature,  (b)  The  test  of  clotting  was  conveni- 
ently made  at  intervals  of  a  half -minute,  and  if  the 
clotting  process  were  hastened  by  higher  tempera- 


FASTER  COAGULATION  BY  ADRENIN  143 

tures,  this  interval  would  become  relatively  less 
exact,  (c)  A  temperature  of  25°  C.  rather  than 
lower  was  selected  because,  as  Dale  and  Laidlaw  ^ 
have  shown,  the  coagulation  time  is  much  slower 
for  a  given  change  in  temperature  below  25° 
than  for  the  same  change  above.  And  with  slow- 
ing of  the  process  the  end  point,  when  the  determin- 
ation depends  on  supporting  a  weight,  is  less  likely 
to  be  sharp,  (d)  The  researches  undertaken  with 
use  of  this  coagulometer  were  concerned  with  fac- 
tors hastening  the  process.  For  that  reason  and 
for  reason  (b),  a  long  rather  than  a  short  coagula- 
tion time  for  normal  conditions  was  desirable. 

3.  Uniformity  in  the  amount  and  hind  of  con- 
tact with  foreign  surface. — The  capacity  of  the  can- 
nulas was  fairly  uniform,  as  stated  above;  the 
amount  received  in  them  was  fairly  constant ;  and 
the  wire  hanging  in  the  blood  presented  approxi- 
mately the  same  surface  in  different  observations. 

A  further  condition  for  insuring  consistent  treat- 
ment of  the  blood  in  different  cases  was  that  of 
making  the  tests  for  coagulation  always  at  the 
same  intervals.  Below  the  writing  point  of  the 
lever  was  set  an  electromagnetic  signal  E,  which 
recorded  half-minutes.  At  the  moment  a  record 
was  made  by  the  signal  (see  first  signal  mark.  Fig. 
25)  the  clip  on  the  artery  was  opened,  the  blood 
taken,  and  the  process  thus  begun.  In  about  20 
seconds  the  cannula  was  suspended  in  the  water 


144 


BODILY   CHANGES 


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FASTER  COAGULATION  BY  ADRENIN  145 

bath  and  the  wire  was  hanging  on  the  lever.  At  the 
next  record  by  the  signal  and  at  every  subsequent 
record  the  vertical  rod  R-  was  pushed  with  the 
index  finger  from  post  P^  to  post  P-  and  allowed  to 
move  back.  This  motion  was  uniform  and  lasted 
about  one  second.  The  check  R^  on  the  long  arm  of 
the  lever  was  thus  raised,  and  as  the  wire  sank  in 
the  blood  the  writing  point  rose,  recording  that 
coagulation  had  not  taken  place  (see  Fig.  25). 

4.  Definite  end  point. — As  soon  as  the  blood  clot- 
ted, the  weight  of  30  milligrams  was  supported,  and 
the  failure  of  the  lever  to  rise  to  the  former  height 
in  the  regular  time  allowed,  recorded  that  the 
change  had  occurred. 

Very  rarely  the  swing  of  the  lever  would  be 
checked  for  a  moment  and  would  then  begin  to 
move  rapidly,  indicating  that  a  strand  of  fibrin  had 
formed  but  not  sufficiently  strong  to  support  the 
weight,  and  that  when  the  strand  broke,  the  weight 
quickly  sank  in  the  blood.  If  this  occurred,  the 
next  record  almost  always  was  the  short  line,  which 
signified  that  the  weight  was  well  supported. 

A  very  slight  strand  of  fibrin  was  able  to  pre- 
vent the  weight  from  dropping,  though  at  different 
times  the  amount  of  support  differed,  as  shown  by 
the  varying  length  of  the  final  lines  (compare  first 
and  last  series.  Fig.  25).  These  variations  are 
probably  a  rough  indication  of  the  degree  of  coagu- 
lation.   In  our  experiments,  however,  the  length  of 


146  BODILY   CHANGES 

the  final  line  was  disregarded,  and  merely  the  fact 
that  the  lever  failed  to  swing  through  its  usual 
distance  was  taken  as  evidence  of  a  clot,  and  the 
consequent  short  record  was  taken  as  the  end  point. 

As  soon  as  this  end  point  was  registered,  the 
tube,  wire  and  cannula  were  lifted  out  of  the  bath ; 
the  cannula  was  then  separated  from  the  tube  and 
pulled  away  from  the  wire.  The  clot  was  thus  dis- 
closed, confirming  the  graphic  record. 

The  method,  at  least  when  used  at  half -minute 
intervals,  did  not  reveal  in  all  instances  the  same 
degree  of  clotting.  Usually,  when  the  process  was 
very  rapid,  the  revealed  clot  was  a  thick  jelly; 
whereas,  when  the  process  was  slow,  a  strand  of 
fibrin  or  at  most  a  small  amount  of  jelly  was  found. 
This  difference  in  the  degree  of  coagulation  intro- 
duced, of  course,  an  element  of  inexactness.  In  our 
experiments,  however,  this  inexactness  was  unfa- 
vorable to  the  result  we  were  seeking  for,  i.  e.,  the 
acceleration  of  the  process — ^because  the  jelly  is  a 
later  stage  than  the  fibrin  strand;  and  since  we 
nevertheless  obtained  good  evidence  of  accelera- 
tion, we  did  not  in  these  experiments  attempt  to 
determine  more  accurately  differences  in  the  stage 
of  the  clotting  process. 

5.  Cleaning  of  apparatus. — After  the  wire  was 
removed  from  the  tube,  the  clot  attached  to  its 
ring-tip  was  carefully  brushed  away  under  cool 
running  water.    Under  the  running  water,  also,  a 


FASTER  COAGULATION  BY  ADRENIN  147 

trimmed  feather  was  introduced  into  the  cannula 
and  the  tube  to  push  out  the  plasticine  and  to  wash 
out  the  blood.  Wire,  cannula  and  tube  were  then 
drojDped  into  a  beaker  receiving  running  hot  water 
(about  80°  C.)  and  there  allowed  to  remain  for 
about  five  minutes.  On  removal  from  this  the 
parts  were  shaken  free  from  water,  passed  through 
95  per  cent  alcohol  and  again  shaken  free,  passed 
through  ether  and  let  dry. 

By  having  a  half-dozen  cannulas  and  wires  of 
standard  size,  it  was  possible  to  save  trouble  by 
cleaning  a  number  at  one  time. 

Not  infrequently  the  first  few  samples  of  blood 
taken  from  an  animal  showed  rapid  or  somewhat 
irregular  rates  of  clotting.  Some  causes  for  these 
initial  variations  will  be  presented  in  following 
pages.  The  fairly  uniform  rate  of  clotting  in  any 
individual  after  the  initial  stage,  varied  in  twenty- 
one  different  animals  from  an  average  of  3  to  an 
average  of  10.6  minutes,  with  a  combined  average 
of  5.9  minutes.  The  conditions  for  these  variations 
among  the  individuals  have  not  been  wholly  deter- 
mined. 

The  Effects  of  Subcutaneous  Injections  of  Adrenin 

The  first  observations  were  of  this  class. 

Oct.  27.  A  cat  weighing  about  3  kilos  was  given 
3  cubic  centimeters  of  adrenin  1 :1,000,  i.e.,  1  milli- 
gram per  kilo,  under  the  skin.    The  animal,  in  this 


148  BODILY   CHANGES 

instance,  was  kept  in  uniform  ether  anesthesia. 
Following  is  a  record  showing  when  blood  was 
taken,  and  the  coagulation  time  in  each  instance: 

2.56— Injection  made  3.2Y— 3.5  minutes 

.59—6      minutes  .44—2 

3,07—5.5         "  .55—2.5 

13—5  "  4.07—3  " 

'.20-6.5         "  -20-2 

Average  5.7  minutes  Average  2.6  minutes 

4.44 — 6      minutes 

5.00—4.5 

5.50—5 

Average  5.2  minutes 

In  this  case  the  coagulation  time  remained  at  its 
usual  level  for  about  20  minutes  after  the  subcu- 
taneous injection.*  Thereafter  for  about  an  hour 
the  coagulation  time  averaged  45  per  cent  of  its 
previous  duration.  And  widely  separated  tests 
made  during  the  following  hour  indicated  that  ap- 
proximately the  initial  rate  of  clotting  had  been  re- 
gained. 

The  rather  long  period  (nearly  30  minutes),  in 
the  case  just  cited,  between  the  injection  and  the 

*  This  period  is  longer  than  is  expected  after  the  subcuta- 
neous injection  of  any  drug.  As  vs^ill  be  shown  later,  strong 
doses  of  adrenin,  if  injected  rapidly,  may  not  at  first  shorten 
the  clotting  process.  Probably  in  some  instances  of  subcu- 
taneous injection  of  these  strong  doses,  the  drug  enters  the 
circulation  more  rapidly  than  in  others  and  in  consequence 
coagulation  is  not  at  first  accelerated. 


FASTER  COAGULATION  BY  ADRENIN  149 

first  appearance  of  rapid  clotting  was  not  the  rule. 
As  the  following-  figures  show,  the  coagulation  time 
may  become  shortened  quite  promptly  after  sub- 
cutaneous injection. 


Oct.  29.  3.30 — 5.5  minutes 

3.53—4     min 

.36—5.5 

4.01—3.5 

.44   Adrenin,    3    cu- 

.08—3.5        " 

bic    centimeters, 

.16^.5         " 

1:1,000,  injected 

.23—5            " 

subcutaneously. 

.30—5.5 

.46 — 5.5  minutes 

In  this  case  nine  minutes  after  the  injection  the 
change  in  the  rate  of  clotting  had  begun,  and  it  con- 
tinued more  rapid  for  the  subsequent  half -hour. 

We  did  not  attempt  to  find  the  minimal  subcu- 
taneous dose  which  would  shorten  clotting.  A 
dose  of  0.01  milligram  per  kilo,  however,  has 
proved  effective,  as  shown  by  the  following  figures : 

Feb.  3.  11.34—10  minutes  .55—10     minutes 

.45—  9        "  12 .06—  7  " 

.50  to  .52  Adrenin,  .14—  4  " 

2.8     cubic     centimeters,  .19 —  5.5 

1 :100,000,  injected  under  .31—  6  " 

skin    of    groin    in    cat  .37 —  7 

weighing  2.8  kilos.  ;45—  9  " 

As  will  be  shown  later,  the  dose  in  this  instance 
was  ten  times  the  minimal  effective  intravenous 
dose.  On  the  basis  of  these  figures,  less  than  a 
milligram  of  adrenin  given  subcutaneously  would 
be  necessary  to  shorten  clotting  to  a  marked  degree 
in  a  man  of  average  weight  (70  kilograms). 


150  BODILY   CHANGES 

Not  many  observations  were  made  by  us  on  the 
effects  of  adrenin  administered  subcutaneously. 
The  amount  reaching  the  vascular  system  and  the 
rate  of  its  entrance  into  the  blood  could  be  so  much 
more  accurately  controlled  by  intravenous  than  by 
subcutaneous  introduction  that  most  of  our  atten- 
tion was  devoted  to  the  latter  method. 

The  Effects  of  Intravenous  Injections 

In  this  procedure  a  glass  cannula  was  fastened 
in  one  of  the  external  jugular  veins  and  filled  with 
the  same  solution  as  that  to  be  injected.  A  short 
rubber  tube  was  attached  and  tightly  clamped  close 
to  the  glass.  Later,  for  the  injection,  the  syrmge 
needle  was  inserted  through  the  rubber  and  into 
the  fluid  in  the  cannula,  the  clip  on  the  vein  was 
removed,  and  the  injection  made. 

The  solutions  employed  intravenously  were 
adrenin  1:10,000,  1:50,000,  and  1:100,000,  in  dis- 
tilled water. 

The  smallest  amount  which  produced  any  change 
in  clotting  time  was  0.1  cubic  centimeter  of  a  dilu- 
tion of  1 :100,000  in  a  cat  weighing  two  kilos,  a  dose 
of  0.0005  railligram  per  kilo.  Four  tests  previous 
to  the  injection  averaged  5  minutes,  and  none  was 
shorter  than  4  minutes.  Immediately  after  the  in- 
jection the  time  was  2  minutes,  but  at  the  next 
test  the  effect  had  disappeared.  Doubling  the 
dose  in  the  same  cat — i.  e.,  giving  0.2  cubic  centi- 


FASTER  COAGULATION  BY  ADRENIN   151 

meter  (0.001  milligram  per  kilo) ^shortened  the 
coagulation  time  for  about  40  minutes : 


Dec.  23. 10  .30 — i      minutes 

10.53—3.5  minutes 

.35-4 

11 .00—1.5 

.41—4 

.05—1.5 

.46  Adrenin,  0.001 

.10—3 

milligram    per 

.15—2 

kilo. 

.20—4 

.4Y — 2.5  minutes 

.26—4.5 

.50—3 

.31—5 

From  10.47,  immediately  after  the  second  injec- 
tion, till  11.20  the  average  time  for  clotting  was  2.5 
minutes,  whereas  both  before  and  after  this  period 
the  time  was  4  minutes  or  longer.  At  11.00  o'clock 
and  11.05,  when  the  end  point  was  reached  in  1.5 
minutes  (  a  reduction  of  63  per  cent),  a  thick  jelly 
was  found  on  examining  the  cannula.  The  changes 
in  clotting  time  in  this  case  are  represented  graph- 
ically in  Fig.  26. 

In  another  case  a  dose  of  0.0005  milligram  per 
kilo  failed  to  produce  any  change,  but  0.001  milli- 
gram per  kilo  (0.28  cubic  centimeter  of  adrenin, 
1 :100,000,  given  a  cat  weighing  2.8  kilos)  brought  a 
sharp  decline  in  the  record,  as  follows : 

Jan.  9.  11 .32 — 6     minutes  11.48 — 5.5  minutes 

.40—6           "  .55—4            " 

.47  Adrenin,  0.001  12.00—5.5          " 

milligram    per  .06 — 7            " 

kilo. 

In  these  instances  the  animals  were  decere- 
brated.   For  decerebrate  cats,  the  least  amount  of 


152 


BODILY   CHANGES 


adrenin,  intravenously,  needed  to  produce  shorten- 
ing of  coagulation  time  is  approximately  0.001 
milligram  per  kilo. 

In  the  above  cases  rapid  clotting  was  manifest 
directly  after  minute  doses.    Larger  doses,  how- 


11111/ 


10:30        :40 


:50      11:00 


:10 


:S0     :30 


Figure  26. — Shortening  of  coagulation 
time  after  injection  of  adrenin,  0.2  cubic 
centimeter,  1:100,000,  (0.001  milligram  per 
kilo),  at  10:46.  In  this  and  following  Fig- 
ures a  scale  for  coagulation  time  is  given  in 
minutes  at  the  left. 

ever,  may  produce  primarily  not  faster  clotting  but 
slower,  and  that  may  be  followed  in  turn  by  a  much 
shorter  coagulation  time.  The  figures  below  pre- 
sent such  an  instance : 


Nov.  25.  2 .36—3     minutes 

3.00—2.5  minutes 

.40—3 

.03—1.5 

.43  Adrenin,       0.5 

05—1.5        " 

cubic  centime- 

.07—2.5 

ter,  1:10,000. 

.10—1.5        « 

.44 — 4     minutes 

.14r-1.5             " 

.49—3.5 

.16—2.5        " 

.53—1.5         " 

19—3           « 

.55—1.5         « 

.23—3           « 

.58—2           « 

.30     3           " 

FASTER  COAGULATION  BY  ADRENIN  153 

This  unexpected  primary  increase  of  coagula- 
tion time,  lasting  at  least  six  minutes,  is  in  strik- 
ing contrast  to  the  later  remarkable  shortening  of 
the  i^rocess  from  3  to  an  average  of  1.7  minutes 
for  more  than  20  minutes  (see  Fig.  27,  A). 

If  a  strong  solution,  i.  e.,  1 :10,000,  is  injected 
rapidly,  the  process  may  be  prolonged  as  above, 
but  not  followed  as  above  by  shortening,  thus : 


Nov.  28.     9  .59—3      minutes 

10 .14—3.5 

minutes 

10 .03—3 

.18—3.5 

.08  Adrenin,     0.5 

.22—3.5 

cubic      centi- 

.26—3 

meter,    1 :10,- 

.29—3 

000. 

.33—3 

.10 — 3      minutes 

There  was  in  this  case  no  decrease  in  coagulation 
time  at  any  test  for  a  half-hour  after  the  injection, 
but  instead  a  lengthening  (see  Fig.  27,  B).  How- 
ell ^  has  reported  the  interesting  observation  that 
repeated  massive  doses  of  adrenin  given  to  dogs 
may  so  greatly  retard  coagulation  that  the  animals 
may  be  said  to  be  hemophilic.  These  two  instances 
show  that  on  coagulation  large  doses  have  the 
contrary  effect  to  small,  just  as  Hoskins  ^°  showed 
was  true  for  intestinal  and  Lyman  and  I  ^^  showed 
was  true  for  arterial  smooth  muscle. 

In  a  few  experiments  the  brain  and  the  cord  to 
midthorax  were  destroyed  through  the  orl)it.  Arti- 
ficial respiration  then  maintained  the  animal  in  uni- 


154 


BODILY    CHANGES 


form  condition.  Under  these  circumstances,  adre- 
nin  intravenously  had  more  lasting  effects  than 
when  given  to  the  usual  decerebrate  animals  with 
intact  cord.  Fig.  28  illustrates  such  a  case.  For 
thirty  minutes  before  injection  the  clotting  time 
averaged  5.4  minutes.  Then,  about  ten  minutes 
after  one  cubic  centimeter  of  adrenin,  1 :50,000,  had 


fi:40 


30     10:00    :10 


:20 


:30 


Figure  27. — A,  Primary  lengthening  followed  by 
shortening  of  the  coagulation  time  when  adrenin,  0.5 
cubic  centimeter  1:10,000  (0.05  milligram),  was  injected 
slowly  at  2 :43.  B,  Lengthening  of  the  coagulation  time 
without  shortening  when  the  same  dose  was  injected 
rapidly  at  10:08. 


been  slowly  injected,  clotting  began  to  quicken; 
during  the  next  twenty  minutes  the  average  was  3.4 
minutes,  and  during  the  following  forty-five  min- 
utes the  average  was  1.9  minutes — only  35  per  cent 
as  long  as  it  had  been  before  the  injection. 

In  another  case  in  which  the  brain  and  upper 
cord  were  similarly  destroyed,  the  clotting  time, 
which  for  a  half -hour  had  averaged  3.9  minutes, 
was  reduced  by  one  cubic  centimeter  of  adrenin, 


FASTER  COAGULATION  BY  ADRENIN  155 

1 :100,000,  to  an  average  for  the  next  hour  and  forty- 
minutes  of  2.3  minutes,  with  1.5  and  3  minutes  as 
extremes.  During  the  first  forty  minutes  of  this 
period  of  one  hour  and  forty  minutes  of  rapid  clot- 
ting all  of  eight  tests  except  two  showed  a  coagula- 
tion time  of  2  minutes  or  less.  The  explanation  of 
this  persistent  rapid  clotting  in  animals  with  spinal 
cord  pithed  is  not  yet  clear. 

As  indicated  in  Figs.  26,  27  and  28,  the  records 
of  coagulation  show  oscillations.  Some  of  these 
ups  and  downs  are,  of  course,  within  the  limits  of 


Figure  28. — Persistent  shortening  of  the  coagulation  time  after 
injecting  (in  an  animal  with  brain  and  upper  cord  pithed)  adrenin, 
1  cubic  centimeter,  1:50,000  (0.02  milligram),  at  11:01-02.  The 
dash  lines  represent  averages. 

error  of  the  method,  but  in  our  experience  they 
have  occurred  so  characteristically  after  injection 
of  adrenin,  and  so  often  have  appeared  in  a  rough 


156  BODILY   CHANGES 

rhythm,  that  they  have  given  the  impression  of  be- 
ing real  accompaniments  of  faster  clotting.  It 
may  be  that  two  factors  are  operating,  one  tending 
to  hasten,  the  other  to  retard  the  process,  and  that 
the  equilibrium  disturbed  by  adrenin  is  recovered 
only  after  interaction  to  and  fro  between  the  two 
factors. 

The  oscillations  in  coagulation  time  after  the  in- 
jections suggest  that  clotting  might  vary  with 
changes  in  blood  pressure,  for  that  also  commonly 
oscillates  after  a  dose  of  adrenin  (see,  e.g..  Fig. 
23).  Simultaneous  recording  of  blood  pressure 
and  determining  of  coagulation  time  have  revealed 
that  each  may  vary  without  corresponding  varia- 
tion in  the  other.  Within  ordinary  limits,  there- 
fore, changes  of  blood  pressure  do  not  change  the 
rate  of  clotting. 

The  Hastening  op  Coagulation  by  Adrenin  Not  a  Direct 
Effect  on  the  Blood 

As  previously  stated,  von  den  Velden  has  con- 
tended that  shortening  of  coagulation  time  by  adre- 
nin is  due  to  exudation  of  tissue  juices  resulting 
from  vasoconstriction.  The  amount  of  adrenin 
which  produces  markedly  faster  clotting  in  the 
cat,  is  approximately  0.001  milligram  per  kilo.  As 
Lyman  and  1 12  showed,  however,  this  amount 
when  injected  slowly,  as  in  the  present  experi- 
ments, results  in  brief  vasodilation  rather  than 


FASTER  COAGULATION  BY  ADRENIN  157 

vasoconstriction.  Von  den  Velden's  explanation 
can  therefore  not  be  applied  to  these  experiments. 
He  has  claimed,  furthermore,  that  adrenin  added 
to  blood  in  vitro  makes  it  clot  more  rapidly,  but, 
as  already  noted,  he  gives  no  account  of  the  condi- 
tions of  his  experiments  and  no  figures.  It  is  im- 
possible, therefore,  to  criticise  them.  His  claim, 
however,  is  contrary  to  Wiggers's  ^^  earlier  ob- 
servations that  blood  with  added  adrenin  coagulat- 
ed no  more  quickly  than  blood  with  an  equal 
amount  of  added  physiological  salt  solution.  Also 
contrary  to  this  claim  are  the  following  two  experi- 
ments: (1)  Ligatures  were  tied  around  the  aorta 
and  inferior  vena  cava  immediately  above  the  dia- 
phragm, and  thus  the  circulation  was  confined  al- 
most completely  to  the  anterior  part  of  the  animal. 
Indeed,  since  the  posterior  part  ceases  to  function 
in  the  absence  of  blood  supply,  the  preparation 
may  be  called  an  "anterior  animal."  When  such  a 
preparation  was  made  and  0.5  cubic  centimeter  of 
adrenin,  1:100,000  (half  the  usual  dose,  because, 
roughly,  half  an  animal),  was  injected  slowly  into 
one  of  the  jugulars,  coagulation  was  not  shortened. 
Whereas  for  a  half-hour  before  the  injection  the 
clotting  time  averaged  4.6  minutes,  for  an  hour 
thereafter  the  average  was  5.3  minutes — a  pro- 
longation which  may  have  been  due,  not  to  any  in- 
fluence of  adrenin,  but  to  failure  of  the  blood  to 
circulate  through  the  intestines  and  liver.^^    In  an- 


158  BODILY   CHANGES 

other  experiment  after  the  gastro-intestinal  canal 
and  liver  had  been  removed  from  the  animal,  the 
average  time  for  coagulation  during  twenty-five 
minutes  before  injecting  adrenin  (0.23  cubic  centi- 
meter, 1 :100,000,  in  an  animal  weighing  originally 
2.3  kilos)  was  5.5  minutes,  and  during  forty  min- 
utes after  the  injection  it  was  6.8  minutes,  with  no 
case  shorter  than  6  minutes.  In  the  absence  of  cir- 
culation through  the  abdominal  viscera,  therefore, 
adrenin  fails  to  shorten  the  clotting  time.  (2)  The 
cannulas  were  filled  with  adrenin,  1:1,000,  and 
emptied  just  before  being  introduced  into  the 
artery.  The  small  amount  of  adrenin  left  on  the 
walls  was  thus  automatically  mixed  with  the  drawn 
blood.  Alternate  observations  with  these  cannulas 
wet  by  adrenin  and  with  the  usual  dry  cannulas 
showed  no  noteworthy  distinction. 

Feb.  19.     2.21 — 6     minutes,  with  usual  cannula 
.30—6.5         «             "  "  " 

.36—6.5         "  "  adrenin     " 

.49—6  "  "  "  " 

.56—7  "  "  usual 

3.04^6  "  "  adrenin     " 

The  results  of  these  experiments  have  made  it 
impossible  for  us  to  concede  either  of  von  den 
Velden's  claims,  i.  e.,  that  clotting  occurs  faster  be- 
cause adrenin  is  added  to  the  blood,  or  because 
adrenin  by  producing  vasoconstriction  causes  tis- 
sues to  exude  coagulant  juices. 

Vosburgh  and  Eichards  found  that  coagulation 


FASTER  COAGULATION  BY  ADRENIN  159 

became  more  rapid  as  the  blood  sugar  increased. 
Conceivably  faster  clotting  might  result  from  this 
higher  percentage  of  blood  sugar.  Against  this 
assumption,  however,  is  the  fact  that  clotting  is 
greatly  accelerated  by  0.001  milligram  adrenin  per 
kilo  of  body  weight,  much  less  than  the  dose 
necessary  to  increase  the  sugar  content  of  the 
blood.^^  And  furthermore,  when  dextrose  (3  cubic 
centimeters  of  a  10  per  cent  solution)  is  added  to 
the  blood  of  an  anterior  animal,  making  the  blood 
sugar  roughly  0.3  per  cent,  the  coagulation  time 
is  not  markedly  reduced.  Adrenin  appears  to  act, 
therefore,  in  some  other  way  than  by  increasing 
blood  sugar. 

Since  adrenin  makes  the  blood  clot  much  faster 
than  normally  in  the  intact  animal,  and  fails  to 
have  this  effect  when  the  circulation  is  confined  to 
the  anterior  animal,  the  inference  is  justified  that 
in  the  small  doses  here  employed  adrenin  produces 
its  remarkable  effects,  not  directly  on  the  blood  it- 
self, not  through  change  in  the  extensive  neuro- 
muscular, bony,  or  surface  tissues  of  the  body,  but 
through  some  organ  in  the  abdomen. 

That  exclusion  of  the  liver  from  the  bodily  econ- 
omy, by  ligature  of  its  vessels  or  by  phosphorus 
poisoning,  will  result  in  great  lengthening  of  the 
coagulation  time  has  been  clearly  shown.  The 
liver,  therefore,  seems  to  furnish  continuously  to 
the  blood  a  factor  in  the  clotting  process  which  is 


160  BODILY   CHANGES 

being  continuously  destroyed  in  the  body.  It  is  not 
unlikely  that  adrenin  makes  the  blood  clot  more 
rapidly  by  stimulating  the  liver  to  discharge  this 
factor  in  greater  abundance.  But  proof  for  this 
suggestion  has  not  yet  been  established. 

EEFERENCES 

1  Vosburgh  and  Eichards :  American  Journal  of  Physi- 
ology, 1903,  ix,  p.  39. 

2  Wiggers :    Archives     of    Internal    Medicine,     1909,     iii, 

p.  152. 

3  Von    den    Velden :    Miinchener    medizinische    Wochen- 

schrift,  1911,  Iviii,  p.  187. 

4  Dale  and  Laidlaw:  Journal  of  Pathology  and  Bacteriol- 
ogy, 1912,  xvi,  p.  362. 

5  Cannon  and  Gray :  American  Journal  of  Physiology, 
1914,  xxxiv,  p.  321. 

«  Cannon  and  Mendenhall :  American  Journal  of  Physi- 
ology, 1914,  xxxiv,  p.  225. 

■^  Addis :  Quarterly  Journal  of  Experimental  Physiology, 
1908,  i,  p.  314. 

^  Dale  and  Laidlaw :  Loc.  cit.,  p.  359. 

^  Howell :  American  Journal  of  Physiology,  1914,  xxxiii, 
p.  xiv. 

^o  Hoskins :  American  Journal  of  Physiology,  1912,  xxix, 
p.  365. 

^^  Cannon  and  Lyman :  American  Journal  of  Physiology, 
1913,  xxxi,  p.  376. 

^2  Cannon  and  Lyman :  Loc.  cit.,  p.  381. 

^^  Wiggers:  Loc.  cit.,  p.  152. 

^*  See  Pawlow :  Archiv  f iir  Physiologie,  1887,  p.  458. 
Bohr:  Centralblatt  fiir  Physiologie,  1888,  ii,  p.  263.  Meek: 
American  Journal  of  Physiology,  1912,  xxx,  p.  173.  Gray 
and  Lunt:  Ihid.,  1914,  xxxiv,  p.  332. 

'^Cannon:  American  Journal  of  Physiology,  1914, 
xxxiii,  p.  396. 


CHAPTER  X 

THE    HASTENING    OF    THE    COAGULATION    OF 
BLOOD    IN   PAIN   AND    GREAT   EMOTION 

In  the  foregoing  chapter  evidence  was  presented 
that  the  intravenous  injection  of  minute  amounts 
of  adrenin  hastens  the  clotting  of  blood.  The 
amounts  used  did  not  vary  much  above  or  below 
the  amounts  discharged  by  the  adrenal  glands  after 
brief  stimulation  of  the  splanchnic  nerves,  as 
found  by  H.  Osgood  in  the  Harvard  Laboratory, 
and  may  therefore  be  regarded  as  physiological. 
Since  injected  adrenin  is  capable  of  shortening  the 
coagulation  time,  may  not  the  increased  secretion 
of  the  adrenals  likewise  have  that  eifect?  The  an- 
swer to  this  question  was  the  object  of  an  investi- 
gation by  W.  L.  Mendenhall  and  myself.^ 

The  blood  was  taken  and  its  coagulation  was  re- 
corded graphically  in  the  manner  already  de- 
scribed. In  some  instances  the  cats  were  etherized, 
in  others  they  were  anesthetized  with  urethane,  or 
were  decerebrated.  The  splanchnic  nerves  always 
were  stimulated  after  being  cut  away  from  connec- 

161 


162  BODILY   CHANGES 

tion  with  the  spinal  cord.  Sometimes  the  nerves 
were  isolated  unilaterally  in  the  abdomen;  some- 
times, in  order  to  avoid  manipulation  of  the  abdom- 
inal viscera,  they  were  isolated  in  the  thorax  and 
stimulated  singly  or  together.  A  tetanizing  cur- 
rent was  used,  barely  perceptible  on  the  tongue 
and  too  weak  to  cause  by  spreading  any  contrac- 
tion of  skeletal  muscles. 

Coagulation  Hastened  by  Splanchnic  Stimulation 

That  splanchnic  stimulation  accelerates  the  clot- 
ting of  blood,  and  that  the  effects  vary  in  different 
animals,  are  facts  illustrated  in  the  following 
cases: 

Oct.  25. — A  cat  was  etherized  and  maintained  in 
uniform  ether  anesthesia.  After  forty  minutes  of 
preliminary  observation  the  left  splanchnic  nerves 
were  stimulated  in  the  abdomen.  Following  are 
the  figures  which  show  the  effects  on  the  coagula- 


tion  time : 

3 .00 — 4     minutes 

.03 — 2.5   minutes 

.07—5.5 

.07—2.5 

.14^-4           " 

.11—3 

.32—4.5         " 

.16—2            " 

.39  to    .40  Stimulation 

.20—1.5         " 

of  left  splanchnic. 

.23—4 

.42 — 5      minutes 

.29—5.5         " 

.49—5 

.40—5.5         " 

.56—2 

.50—5            " 

4.00—1            « 

FASTER  COAGULATION  IN  EMOTION  163 


In  this  instance  at  least  ten  minutes  elapsed  be- 
tween the  end  of  stimulation  and  the  beginning  of 
faster  clotting.  The  period  of  faster  clotting,  how- 
ever, lasted  for  about  a  half-hour,  during  which  the 
coagulation  time  averaged  2.1  minutes,  only  forty- 
three  per  cent  of  the  previous  average  of  4.8  min- 
utes.   It  is  noteworthy  that  the  curve  (see  Fig.  29), 


6 

1       1                             1 

B 

4 
S 

1 

r-\        . 

2 

1 

-■  ^ ,- 

S:SO 


:40 


:50 


4:00 


t20 


:30 


:40 


Figure  29. — Shortening  of  coagulation 
time  after  stimulation  of  the  left  sjjlanchnic 
nerves,  3:39-:40. 

while  lower,  shows  oscillations  not  unlike  those 
which  follow  injection  of  adrenin  (see  p.  155). 

The  primary  delay  of  the  effect  is  not  always, 
indeed  it  is  not  commonly,  present : 

Nov.  6. — A  cat  was  anesthetized  (1.40  p.m.) 
with  urethane,  and  later  (3.05)  its  brain  was 
pithed.  The  following  observations  on  the  coagu- 
lation time  show  the  prompt  effect  of  splanchnic 
stimulation : 


164  BODILY   CHANGES 

3  ^36 — 7      minutes 

.46—6 
4 .02  to  .05  Stimulation  of  left  splanchnic  in  abdomen. 

.08 — 4     minutes 

.10—3 

.18—3.5         " 

.23—6.5         " 

In  Fig.  30  is  presented  the  original  record  of  the 
shortening  of  the  coagulation  after  stimulation  of 
the  left  splanchnic  nerve  (Nov.  8)  in  a  cat  with 
brain  pithed. 

In  the  foregoing  instances  the  coagulation  time 
was  reduced  after  splanchnic  stimulation  to  less 
than  half  what  it  was  before.  The  reduction  was 
not  always  so  pronounced. 

Nov.  7. — A  cat*  maintained  in  uniform  ether 
anesthesia  with  artificial  respiration  had  the  fol- 
lowing changes  in  the  clotting  time  of  its  blood  as 
the  result  of  stimulating  the  left  splanchnic  nerve 
in  the  thorax : 


3 .40 — 5     minutes 

4.06 — 3.5  minutes 

.45—5 

.11^            " 

.51—5.5         " 

.16—3.5 

.58  to  4.00  Stimulation  of 

.21—4            " 

left     splanch- 

.26—4.5 

nic. 

.31—5            " 

4.01 — 4.5  minutes 

.36—6.5        " 

In  this  case  the  average  for  about  fifteen  minutes 
before  stimulation  was  slightly  over  five  minutes, 

*  This  animal  had  just  passed  through  a  period  of  excite- 
ment with  rapid  clotting. 


FASTER  COAGULATION  IN  EMOTION  165 


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166  BODILY    CHANGES 

and  for  twenty-five  minutes  thereafter  it  was  four 
minutes. 

In  all  cases  thus  far  the  period  of  shortened 
coagulation  lasted  from  ten  to  thirty  minutes.  In 
other  cases,  however,  the  effect  was  seen  only  in  a 
single  observation.  If  this  had  occurred  only  once 
after  splanchnic  stimulation,  it  might  be  attributed 
to  accident,  but  it  was  not  an  infrequent  result, 
e.  g. : 

Oct.  28. — A  cat  was  etherized  and  decerebrated, 
and  the  splanchnic  nerves  were  isolated  in  the 
thorax.  Following  are  two  instances  of  brief  short- 
ening of  coagulation  after  splanchnic  stimulation : 

3.36 — i.5  minutes  4.07 — 4.5  minutes 

.42—4.5         "  .12—5.5         " 

.47  to  .49  Splanchnic  stim-  .  19  to  .22  Splanchnic  stim- 
ulation, ulation. 

.51 — i.5  minutes  ,23 — 3.5  minutes 

.57—2  "  .27—4  " 

4.01—4  «  .33—5  " 

In  the  foregoing  instance  it  is  noteworthy  that 
the  degree  of  acceleration  is  not  so  great  after  the 
second  stimulation  of  the  splanchnics  as  it  was 
after  the  first.  This  reduction  of  effect  as  the 
nerves  were  repeatedly  stimulated  was  frequently 
noted.  The  following  case  presents  another  illus- 
tration : 

Nov.  12.— A  cat  was  etherized  (2.35  p.m.)  and 
the  medulla  was  punctured  (piqiire)  at  3.12.    The 


FASTER  COAGULATION  IN  EMOTION  1G7 

operation  was  without  effect.  The  loss  or  lessen- 
ing of  effectiveness  on  second  stimulation  of  the 
left  splanchnic  nerves  is  to  be  compared  with  the 
persistence  of  effectiveness  on  the  right  side: 

3.40 — 4.5  minutes  4.34 — 4  minutes 

,45 — 4.5         "  .39 — 4         " 

.54  to  .56  Stimulation  of  .44 — 4        " 

left    splanchnic  .48 — 4         " 

in   abdomen.  .55  to  .57  Stimulation  of 
4.00—3      minutes  right     splanch- 

.05—2  "  nic. 

.10—5.5         "  .59—3      minutes 

.16—5  "  5.02—2.5         " 

.22  to  .27  Stimulation  of  .07—3  " 

left   splanchnic  .11 — 3  " 

in  abdomen.  .15 — 5.5         " 

.30 — 4      minutes  .22—5.5         " 

The  experiments  above  recorded  show  that 
stimulation  of  the  splanchnic  nerves  results  imme- 
diately, or  after  a  brief  j^eriod,  in  a  shortening  of 
the  coagulation  time  of  the  blood — an  effect  which 
in  different  animals  varies  in  duration  and  intens- 
ity, and  diminishes  as  the  stimulation  is  repeated. 
The  next  question  was  whether  this  effect  is  pro- 
duced through  the  adrenal  glands. 

CoAGUL.\TioN  Not  Hastknkd  by  Splanchnic  Stimulation  if 
THE  Adrenal  Glands  are  Absent 

The  manner  in  which  splanchnic  stimulation  pro- 
duces its  effects  is  indicated  in  the  following  ex- 
periments : 

Nov.  28. — A  cat  was  etherized,  and  through  the 


168  BODILY   CHANGES 

orbit  the  central  nervous  system  was  destroyed  to 
the  midthorax.  The  blood  vessels  of  the  left  adre- 
nal gland  were  then  quickly  tied  and  the  gland 
removed.  The  readings  for  a  half  hour  before  the 
left  splanchnic  nerve  was  stimulated  averaged 
seven  minutes,  then — 

4.38  to  .40  Stimulation  of  left  splanchnic  (glandless). 

.42 — 7  minutes 

.50—7 
5.02  to  .04  Stimulation  of  right  splanchnic. 

.06 — 4  minutes 

.10—7 

.18—7 

.26—7 

Dec.  4. — A  cat  was  etherized  and  pithed  through 
the  orbit  to  the  neck  region.  The  right  and  left 
splanchnic  nerves  were  tied  and  cut  in  the  thorax. 
The  left  adrenal  gland  was  then  carefully  removed. 
These  operations  consumed  about  a  half-hour.  The 
following  records  show  the  effect  of  stimulating 
the  left  and  right  splanchnic  nerves : 


4.10 — 5      minutes 

5.00—2.5 

minutes 

.16-4.5 

.14—6 

i( 

.25  to  .28  Stimulation  of 

.23  to  .25  Stimulation  of 

left  splanchnic 

right    splanch- 

(glandless). 

nic. 

.30 — 4.5  minutes 

.26—6 

minutes 

.35—4.5         " 

.33—4.5 

u 

.40—7.5 

.38—3.5 

<i 

.49  to  .51  Stimulation  of 

.43—4.5 

ce 

right    splanch- 

.49—5 

(( 

nic. 

.55—6 

(C 

.55 — 4.5  minutes 

FASTER  COAGULATION  IN  EMOTION  169 

The  results  in  this  experiment  are  represented 
graphically  in  Fig.  31. 


4:10      :20  :30  :40  :50         5;00         :10  :20  :30  :40  :50 

Figure  31. — Results  of  stimulating  the  left  splanchnic  nerves, 
4:25-:28,  after  removal  of  the  left  adrenal  gland;  and  of  stimu- 
lating the  right  splanchnic  nerves,  4:49-:51  and  5:23-:25,  with 
right  adrenal  gland  present. 


Elliott's  evidence  that  in  the  cat  the  splanchnic 
innervation  of  the  adrenals  is  not  crossed  has  al- 
ready been  mentioned.  If  the  gland  is  removed  on 
one  side,  therefore,  stimulation  of  the  nerves  on 
that  side  causes  no  discharge  from  the  opposite 
gland.  As  the  above  experiments  clearly  show, 
splanchnic  stimulation  on  the  glandless  side  results 
in  no  shortening  of  the  coagulation  time;  whereas, 
in  the  same  animals,  stimulation  of  the  nerves  on 


170  BODILY   CHANGES 

the  other  side   (still  connected  with  the  adrenal 
gland)  produces  a  sharp  hastening  of  the  clotting 

process. 

The  splanchnics  innervate  the  intestines  and 
liver  even  though  the  adrenal  gland  is  removed. 
The  foregoing  experiments  indicate  that  the  nerve 
impulses  delivered  to  these  organs  do  not  influence 
them  in  any  direct  manner  to  accelerate  the  speed 
of  coagulation.  Indeed,  in  one  of  the  experiments 
(Dec.  4,  see  Fig.  31)  a  high  reading  about  ten  min- 
utes after  splanchnic  stimulation  on  the  glandless 
side  suggests  the  possibility  of  an  opposite  effect. 
Direct  stimulation  of  the  hepatic  nerves  on  one 
occasion  was  followed  by  a  change  of  the  clotting 
time  from  4.5,  5,  4.5, 4.5  minutes  during  twenty-five 
minutes  before  stimulation  to  4.5,  7,  and  6  minutes 
during  twenty  minutes  after  stimulation. 

Since  with  the  adrenals  present  stimulation  of 
hepatic  nerves  induces  alteration  of  glycogen  in  the 
liver  and  quick  increase  of  blood  sugar,^  just  as 
splanchnic  stimulation  does,  the  failure  of  the 
blood  to  clot  faster  after  stimulation  of  the  hepatic 
nerves  confirms  the  evidence  already  offered  that 
faster  clotting  when  adrenin  is  increased  in  the 
blood  is  not  due  to  a  larger  amount  of  sugar  pres- 
ent (see  p.  159). 

The  liver  and  intestines  cannot  be  made  to 
shorten  clotting  time  by  stimulation  of  their 
nerves, but, as  has  already  been  shown  (see  p.  157), 


FASTER  COAGULATION  IN  EMOTION  171 

neither  can  adrenin  act  by  itself  to  hasten  the  clot- 
ting process.  Apparently  the  effect  is  produced  by 
cooperation  between  the  adrenals  and  the  liver 
(and  possibly  also  the  intestines) .  Somewhat  simi- 
lar cooperation  is  noted  in  the  organization  of 
sugar  metabolism;  splanchnic  stimulation  in  the 
absence  of  the  adrenal  glands  does  not  increase 
blood  sugar,^  and  in  the  absence  of  the  liver  adre- 
nin is  without  influence.* 

The  variations  of  effect  noted  after  splanchnic 
stimulation  can  be  accounted  for  by  variations 
in  the  adrenin  content  of  the  glands.  Elliott  ^ 
found,  as  previously  stated,  that  animals  newly 
brought  into  strange  surroundings  may  have  a  con- 
siderably reduced  amount  of  adrenin  in  their  adre- 
nals. The  animals  used  in  our  experiments  had 
been  for  varying  lengths  of  time  in  an  animal 
house  in  which  barking  dogs  were  also  kept,  and 
were  therefore  subject  to  influences  which  would  be 
likely  to  discharge  the  glands. 

The  evidence  that  stimulation  of  splanchnic 
nerves,  with  accompanying  increase  of  adrenal 
secretion,  results  in  more  rapid  clotting  of  blood  is 
especially  interesting  in  relation  to  the  experiments 
previously  described,  which  showed  that  in  pain 
and  emotional  excitement  there  is  an  increased 
secretion  of  adrenin  into  the  blood.  Does  the  adre- 
nin thus  liberated  have  any  effect  on  the  rate  of 
coagulation  ?    The  observations  here  recorded  were 


172 


BODILY   CHANGES 


made  in  order  to  obtain  an  answer  to  that  ques- 
tion. 

Coagulation  Hastened  by  "Painful"  Stimulation 

In  the  experiments   on  the  action  of   stimuli 

which  in  the  unanesthetized  animal  would  cause 

pain,  it  will  be  recalled  that  f  aradic  stimulation  of 

a  large  nerve  trunk  (the  stump  of  the  cut  sciatic) 


1 

7 

<l  

■   1 

1   1 

■   /- 

s  — 

3| 

2  

'"V 

i 

1/  - 

1 

m 

1 

*               — 

1  I 

;50       5:00 


:10 


:30 


Figure  32. — Three  shortenings  of  coagulation  time  after  stimu- 
lation of  the  left  sciatic  nerve,  at  4:23-:25,  at  4:45-:50  (stronger), 
and  at  5:15-:17. 

and  operation  under  light  anesthesia  were  the 
methods  used  to  affect  the  afferent  nerves.  El- 
liott^ found  that  repeated  excitation  of  the  sci- 
atic nerve  was  especially  efficient  in  exhausting  the 
adrenal  glands  of  their  adrenin  content,  and  also 


FASTER  COAGULATION  IN  EMOTION  173 

that  this  reflex  persisted  after  removal  of  the  cere- 
bral hemispheres.  It  was  to  be  expected,  there- 
fore, that  with  well-stored  glands,  sciatic  stimula- 
tion, even  in  the  decerebrate  animal,  would  call 
forth  an  amount  of  adrenal  secretion  which  would 
decidedly  hasten  clotting.  The  following  case  il- 
lustrates such  a  result : 

Dec.  12. — A  cat  was  anesthetized  with  ether  at 
3.45  and  the  left  sciatic  nerve  was  bared.  Decere- 
bration  was  completed  at  3.57.  The  clotting  time 
of  the  blood  began  to  be  tested  six  minutes  later: 


4 .03 — i     minutes 

4 .53—2.5 

minutes 

.08—3.5 

.57—7 

a 

.13—3.5 

5 .06—7.5 

a 

.18—4.5 

.15  to  .1' 

J  Stimulation  of 

.23  to  .25   Stimulation 

of 

left  sciatic. 

left   sciatic. 

5.17—4 

minutes 

4.26—2.5  minutes 

.22—4.5 

(( 

.29—3.5 

.27—5.5 

<c 

.34^-4 

.36—5.5 

(( 

.40—5 

.46—7 

(( 

.45  to  .50  Stimulation 

of 

left  sciatic. 

The  results  obtained  in  this  case,  which  were 
similar  to  results  in  other  cases,  are  represented 
graphically  in  Fig.  32.  The  coagulation  time 
was  becoming  gradually  more  prolonged,  but 
each  excitation  of  the  sciatic  nerve  was  followed 
by  a  marked  shortening.  The  strength  of  stimu- 
lation was  not  determined  with  exactness,  but  it 


174  BODILY   CHANGES 

is  worthy  of  note  that  the  current  used  in  the 
first  and  the  third  stimulations  was  weaker  than 
could  be  felt  on  the  tongue,  whereas  that  used  in 
the  second  was  considerably  stronger,  though  it 
did  not  produce  reflex  spasms. 

Mere  tying  of  the  nerve  is  capable  of  producing 
a  marked  shortening  of  coagulation,  as  the  follow- 
ing figures  show : 

Oct.  21. — 10.57  cat  under  ether,  and  urethane 
given : 

11.11 — 8.5  minutes 

.23—8.5 

.32  to  .35  Left  sciatic  bared  and  tied. 

.37 — 1.5  minutes 

.41—5.5 

.50—7 
12.02—8.5        " 

Stimulation  of  the  crural  nerve  had  similar 
effects,  reducing  the  clotting  time  in  one  instance 
from  a  succession  of  3,  3,  and  3.5  minutes  to  1.5 
minutes  shortly  after  the  application  of  the  cur- 
rent, with  a  return  to  3.5  minutes  at  the  next  test. 

Operative  procedures  performed  under  light 
anaesthesia  (i.  e.,  with  the  more  persistent  reflexes 
still  present),  or  reduction  of  anesthesia  soon  after 
operation,  resulted  in  a  remarkable  shortening  of 
the  coagulation  time : 

Nov.  8. — A  cat  was  etherized  and  tracheoto- 
mized.  The  abdomen  was  then  opened  and  a  liga- 
ture was  drawn  around  the  hepatic  nerves.    The 


FASTER  COAGULATION  IN  EMOTION  175 

operation  was  completed  at  2.25.  At  2.50  the 
etherization  became  light  and  the  rate  of  clotting 
began  to  be  faster : 

2.50 — 6      minutes  3.15 — 3.5  minutes 

3.00—5.5         "  .20—4.5         " 

.10—3.5         "  .30—7.5         " 

Nov.  11, — A  female  cat,  very  quiet,  was  placed 
in  the  holder  at  1.55!  The  animal  was  not  excited. 
At  2.10  etherization  was  begun;  the  animal  was 
then  tracheotomized,  and  the  femoral  artery  was 
exposed. 

2.21 — 4.5  minutes 

.26 — 4.5        "        Anesthesia  lessened. 

.32—3.5        "  "  light. 

.35  Abdomen  opened. 

.47 — 1.5  minutes. 

.52—1 

.55  Ligature  passed  around  hepatic  nerves. 

.57 — 1.5  minutes.     Anesthesia  light;  corneal  reflex  present. 
3.02—3 

.07 — 3  "         Some  hepatic  nerves  cut. 

.12 — 4.5         "         Rest  of  hepatic  nerves  cut. 

.22—5 

The  results  of  this  experiment  are  shown  graph- 
ically in  Fig.  33. 

Nov.  13. — A  cat  was  etherized  at  1.55,  tracheoto- 
mized, and  the  femoral  artery  laid  bare.  As  soon 
as  these  preparations  were  completed,  the  ether 
was  removed  and  anesthesia  became  light.  The 
blood  clotted  thus : 


176 


BODILY   CHANGES 


2.08—6 

minutes 

.15—4 

iC 

Anesthesia  light. 

.20—2 

l( 

.24—1 

u 

Etherization  begun  again 

.27—2.5 

(C 

.30     3.5 

i( 

.85—5.5 

(( 

.50—5.5 


In  the  foregoing  and  in  other  similar  instances, 
a  condition  of  surgical  injury,  whether  just  made 


Figure  33. — Shortening  of  coagulation 
time  during  an  operation  under  light  anes- 
thesia. At  2 .35  the  abdomen  was  opened, 
at  2:55  a  hgature  was  passed  around  the 
hepatic  nerves. 

or  being  made,  was  accompanied  by  more  rapid 
clotting  of  blood  when  the  degree  of  anesthesia  was 
lessened.  This  condition  was  one  which,  if  allowed 
to  go  further  in  the  same  direction,  would  result 
in  pain.  Both  direct  electrical  stimulation  and  also 
surgical  operation  of  a  nature  to  give  pain  in  the 
unanesthetized  animal  result,  therefore,  in  faster 
clotting. 


FASTER  COAGULATION  IN  EMOTION  177 

It  is  worthy  of  note  that  after  decerebration  clot- 
ting apparently  occurred  no  faster  because  the  ab- 
domen had  been  opened,  although  in  the  decere- 
brate state  etherization  was  suspended.  The 
mechanism  for  reflex  control  of  the  adrenals  may 
not  be  higher  than  the  corpora  quadrigemina,  as 
Elliott  has  shown,  but  the  discharge  from  the 
glands  seems  to  be  more  certain  to  occur  when  the 
cerebrum  is  present  and  is  permitted  even  slightly 
to  operate. 

Coagulation  Hastened  in  Emotional  Excitement  ~"  I 

The  evidence  for  emotional  secretion  of  the  adre-  ^_ 
nal  glands  has  already  been  presented.  As  was 
noted  in  my  earlier  observations  on  the  motions  of 
the  alimentary  canal  (see  p.  14),  cats  differ  widely 
in  their  emotional  reaction  to  being  bound ;  some, 
especially  young  males,  become  furious ;  others, 
especially  elderly  females,  take  the  experience 
quite  calmly.  This  difference  of  attitude  was  used 
with  positive  results,  the  reader  will  recall,  in  the 
experiments  on  emotional  glycosuria ;  there  seemed 
a  possibility  likewise  of  using  it  to  test  the  effect 
of  emotions  on  blood  clotting.  To  plan  formal  ex- 
periments for  that  purpose  was  not  necessary,  be- 
cause in  the  ordinary  course  of  the  researches  here . 
reported,  the  difference  in  effects  on  the  blood  be- 
tween the  violent  rage  of  vigorous  young  males  and 
the  quiet  complacency  of  old  females  was  early 


178 


BODILY    CHANGES 


noted.  Indeed,  the  rapid  clotting  which  accom- 
panied excitement  not  infrequently  made  necessary 
an  annoying  wait  till  slower  clotting  would  permit 
the  use  of  experimental  methods  for  shortening  the 
process. 

The  animals  used  on  November  11  and  13  (see 
pp.  175,  176)  are  examples  of  calm  acceptance  of 
being  placed  on  the  holder;  and  furthermore,  these 
animals  were  anesthetized  without  much  dis- 
turbance. As  the  figures  indicate,  the  clotting 
from  the  first  occurred  at  about  the  average  rate. 

In  sharp  contrast  to  these  figures  are  those  ob- 
tained when  a  vigorous  animal  is  angered : 

Oct.  30. — A  very  vigorous  cat  was  placed  on  the 
holder  at  9.08.  It  at  once  became  stormy,  snarling, 
hissing,  biting,  and  lashing  its  big  tail.  At  9.12 
etherizing  was  begun  and  that  intensified  the  ex- 
citement. By  9.15  the  femoral  artery  was  tied. 
The  clotting  time  of  the  blood  for  an  hour  after  the 
ether  was  first  given  was  as  follows : 


9.18—0.5     minute                              9.43—1 

minute 

.19—1 

.45—0.5 

a 

.22—1 

.49—0.5 

a 

.24—1 

.52—0.5 

u 

.26—1 

.54—0.5 

a 

.28—1.5          ' 

.57—1 

a 

.31—1 

'                                 10.00—0.5 

u 

.33—0.5 

'                                     .02—0.5 

a 

.35—0.5 

.06—1 

a 

.38—0.5 

'                                     .09—0.5 

a 

.39—0.5          ' 

.11—0.5 

a 

.41—1 

.13—1 

a 

FASTER  COAGULATION  IN  EMOTION  179 

Twenty-four  observations  made  during  the  hour 
showed  that  the  clotting  time  in  this  enraged  ani- 
mal averaged  three-fourths  of  a  minute  and  was 
never  longer  than  a  minute  and  a  half.  The  clots 
were  invariably  a  solid  jelly.  The  persistence  of 
the  rapid  clotting  for  so  long  a  period  after  anes- 
thesia was  started  may  have  been  in  part  due  to 
continued,  rather  light,  etherization,  for  Elliott  '^ 
found  that  etherization  itself  could  reduce  the 
adrenin  content  of  the  adrenal  glands. 

The  shortened  clotting  did  not  always  persist  so 
long  as  in  the  foregoing  instance.  The  brief  period 
of  faster  clotting  illustrated  in  the  following  case 
was  typical  of  many : 

Nov.  18. — A  cat  that  had  been  in  stock  for  some 
time  was  placed  on  the  holder  at  2.13,  and  was  at 
once  enraged.  Two  minutes  later  etherization  was 
started.  The  hairs  on  the  tail  were  erect.  The 
clotting  was  as  follows : 

2.25 — 1      minute.  2.31 — 4.5  minutes 

.27—0.5        "  .37—3.5        " 

.28—2  "  .47—4.5        " 

It  seems  probable  that  in  this  case  just  as  in 
some  of  the  cases  in  which  the  splanchnic  nerves 
were  stimulated  (see  p.  166),  the  adrenals  had  been 
well-nigh  exhausted  because  of  the  cat's  being 
caged  near  dogs,  and  that  the  emotional  flare-up 
practically  discharged  the  glands,  for  repeated  at- 


180  BODILY   CHANGES 

tempts  later  to  reproduce  the  initial  rapid  clotting 
by  stimulation  of  the  splanchnic  nerves  were  with- 
out result. 

Evidence  presented  in  previous  chapters  makes 
wholly  probable  the  correctness  of  the  inference 
that  the  faster  coagulation  which  follows  emotional 
excitement  is  due  to  adrenal  discharge  from 
splanchnic  stimulation.  In  this  relation  the  effect 
of  severance  of  the  splanchnics  on  emotional  accel- 
eration of  the  clotting  process  is  of  interest.  The 
following  cases  are  illustrative : 

Oct,  29. — A  cat  was  left  on  the  holder  for  ten 
minutes  while  the  femoral  artery  was  uncovered 
under  local  anesthesia.  The  blood  removed  was 
clotted  in  a  half -minute.  The  animal  was  much 
excited.  It  was  now  quickly  etherized  and  the 
brain  pithed  forward  from  the  neck.  The  tests 
resulted  as  follows : 

10.51 — 1      minute. 

.53—0.5 

.55—0.5         " 

.57—0.5         " 
11.07  Cut  left  splanchnic. 

.12     "     right  splanchnic. 

.21 — 3.5  minutes. 

.26—3.5         " 

The  original  record  of  this  case  is  given  in  Fig. 
34. 

Nov.  5.— A  cat  was  etherized  at  2.35.  At  2.39 
artificial  respiration  by  tracheal  cannula  was  be- 


FASTER  COAGULATION  IN  EMOTION  181 

gun,  the  air  passing  through  an  ether  bottle.    The 

clotting  occurred  thus: 

2.53 — 1.5  minutes 

.57—1.5 

3.05—1.5 

.15—1.5 

.25  Both  splanchnics  cut  and  tied  in  thorax. 

.35 — i.5  minutes 

.55—4.5 

Nov.  7. — A  cat  was  etherized  at  1.55  under  ex- 

citement and  with  tail  hairs  erect.    At  2.13  the  ani- 

H^^^^KM^^^^^Sf^^^^H^^H^^HIHHH 

;J.    -  •  -    -     lii      ,    n 

1     1    1-  >--t — r— t— 1 — t— 1 — 1— 1 — t— t— t w — t— t — (— t— 1 — 1 — 1 — 1 — 1 — 1     1    1    1 — h— 1 — 1— t- 

10:61 


10:53 


10:55         10:57 


11:21 


Figure  34. — About  two- thirds  original  size.  Record  of  rapid 
clotting  (less  than  a  half-minute)  after  emotional  excitement.  At 
11:07  the  left,  at  11:12  the  right  splanchnic  nerves  were  cut;  the 
clotting  then  required  3:5  minutes.  The  marks  below  the  time 
record  indicate  the  moments  when  the  samples  were  drawn. 

mal  was  showing  reflexes.    The  figures  show  the 
course  of  the  exijeriment : 


2.15—1.5 

minutes 

.21—1 

.26—1 

.31—1 

.36—1 

.41—1 

.46—2 

.51—2 

3.06—2 

3.11 — 2.5  minutes 
.26  Cut    left    splanchnic    in 

thorax. 
.35  Cut  ri^ht  splanchnic  in 

thorax. 
.40 — 5      minutes 
.45—5  " 

.51—5.5         " 


182  BODILY    CHANGES 

In  this  instance  the  subsequent  stimulation  of 
the  splanchnic  nerves  resulted  again  in  faster  clot- 
ting—a reduction  from  5.5  minutes  to  3.5  minutes 
(see  experiment  Nov.  7,  p.  164).  The  results  from 
this  experiment  are  expressed  graphically  in  Fig. 
35. 


1      1 

5  - 

1              t 

1              1              1              1              1 

4  ~ 

- 

8  ^ 

- 

2  — 

/ 

1              1 

/ 

1             \ 

1             I              1              1              1 

2:10      :20         :30         :40         :50         S:00        :10  :20  :30         :40         :50 

Figure  35. — Rapid  clotting  after  emotional  excitement,  with 
slowing  of  the  process  when  the  splanchnic  nerves  were  cut  in  the 
thorax  (the  left  at  3:26,  the  right  at  3:35). 

The  data  presented  in  this  chapter  show  that 
such  stimulation  as  in  the  unanesthetized  animal 
would  cause  pain,  and  also  such  emotions  as  fear 
and  rage,  are  capable  of  greatly  shortening  the 
coagulation  time  of  blood.  These  results  are  quite 
in  harmony  with  the  evidence  previously  offered 
that  injected  adrenin  and  secretion  from  the  adre- 
nal glands  induced  by  splanchnic  stimulation 
hasten  clotting,  for  painful  stimulation  and  emo- 


FASTER  COAGULATION  IN  EMOTION  183 

tional  excitement  also  evoke  activity  of  the  adre- 
nals. Here,  then,  is  another  fundamental  change 
in  the  body,  a  change  tending  to  the  conservation 
of  its  most  important  fluid,  wrought  through  the 
adrenal  glands  in  times  of  great  perturbation. 
This  bodily  change  and  the  others  which  occur 
under  the  same  circumstances  are  next  to  be  ex- 
amined with  reference  to  their  significance. 

REFERENCES 

^  Cannon  and  Mendenhall :  American  Journal  of  Physi- 
ology, 1914,  xxxiv,  p.  251. 

2  Macleod :  Diabetes :  its  Pathological  Physiology,  Lon- 
don, 1913,  pp.  68-72. 

^  Gautrelet  and  Thomas:  Comptes  Rendus,  Societe  de 
Biologie,  1909,  Ixvii,  p.  233. 

*  Bang :  Der  Blutzucker,  Wiesbaden,  1913,  p.  87. 

5  Elliott:  Journal  of  Physiology,  1912,  xliv,  p.  379. 

6  Elliott :  Loc.  cit.,  pp.  406,  407. 

7  Elliott :  Loc.  cit.,  p.  388. 


CHAPTEE  XI 

THE    UTILITY    OF    THE    BODILY    CHANGES    IN 
PAIN  AND   GREAT   EMOTION 

We  now  turn  from  a  consideration  of  the  data 
secured  in  our  experiments  to  an  interpretation  of 
the  data.  One  of  the  most  important  lessons  of  ex- 
perience is  learning  to  distinguish  between  the 
facts  of  observation  and  the  inferences  drawn  from 
those  facts.  The  facts  may  remain  unquestioned; 
the  explanation,  however,  may  be  changed  by  addi- 
tional facts  or  through  the  influence  of  more  ex- 
tensive views.  Having  given  this  warning,  I  pro- 
pose to  discuss  the  bearings  of  the  results  reported 
in  the  earlier  chapters. 

Our  inquiry  thus  far  has  revealed  that  the 
adrenin  secreted  by  the  adrenal  glands  in  times  of 
stress  has  all  the  effects  in  the  body  that  are  pro- 
duced by  injected  adrenin.  It  plays  an  essential 
role  in  calling  forth  stored  carbohydrate  from  the 
liver,  thus  flooding  the  blood  with  sugar;  it  helps 
in  distributing  the  blood  to  the  heart,  lungs,  central 
nervous  system  and  limbs,  while  taking  it  away 

184 


UTILITY   OF   BODILY   CHANGES     185 

from  the  inhibited  organs  of  the  abdomen;  it 
quickly  abolishes  the  effects  of  muscular  fatigue ; 
and  it  renders  the  blood  more  rapidly  coagulable. 
These  remarkable  facts  are,  furthermore,  asso- 
ciated with  some  of  the  most  primitive  experiences 
in  the  life  of  higher  organisms,  experiences  com- 
mon to  all,  both  man  and  beast — the  elemental 
experiences  of  pain  and  fear  and  rage  that  come 
suddenly  in  critical  emergencies.  What  is  the  sig- 
nificance of  these  profound  bodily  alterations! 
What  are  the  emergency  functions  of  secreted 
adrenin  ? 

The  Reflex  Nature  of  Bodily  Responses  in  Pain  and 

THE  Major  Emotions,  and  the  Useful  Character 

OF  Reflexes 

The  most  significant  feature  of  these  bodily  re- 
actions in  pain  and  in  the  presence  of  emotion- 
provoking  objects  is  that  they  are  of  the  nature 
of  reflexes — they  are  not  willed  movements,  indeed 
they  are  often  distressingly  beyond  the  control 
of  the  will.  The  pattern  of  the  reaction,  in  these 
as  in  other  reflexes,  is  deeply  inwrought  in  the 
workings  of  the  nervous  system,  and  when  the 
appropriate  occasion  arises,  typical  organic  re- 
sponses are  evoked  through  inherent  automatisms. 

It  has  long  been  recognized  that  the  most  char- 
acteristic feature  of  reflexes  is  their  "purposive" 
nature,  or  their  utility  either  in  preserving  the 


186  BODILY   CHANGES 

welfare  of  the  organism  or  in  safeguarding  it 
against  injury.  The  reflexes  of  sucking,  swal- 
lowing, vomiting  and  coughing,  for  instance,  need 
only  to  be  mentioned  to  indicate  the  variety  of 
ways  in  which  reflexes  favor  the  continuance  of 
existence.  When,  therefore,  these  automatic  re- 
sponses accompanying  pain  and  fear  and  rage— 
the  increased  discharge  of  adrenin  and  sugar— are 
under  consideration,  it  is  reasonable  to  inquire 
first  as  to  their  utility. 

Numerous  ingenious  suggestions  have  been  of- 
fered to  account  for  the  more  obvious  changes 
accompanying  emotional  states — as,  for  example, 
the  terrifying  aspect  produced  by  the  bristling  of 
the  hair  and  the  uncovering  of  the  teeth  in  an 
access  of  rage.^  The  most  widely  applicable  ex- 
planation proposed  for  these  spontaneous  reac- 
tions is  that  during  the  long  course  of  racial 
experience  they  have  been  developed  for  quick 
service  in  the  struggle  for  existence.  Earlier 
writers  on  organic  evolution  pointed  out  the  antici- 
patory character  of  these  responses.  According  to 
Spencer,^  "Fear,  when  strong,  expresses  itself 
in  cries,  in  efforts  to  hide  or  escape,  in  palpitations 
and  tremblings ;  and  these  are  just  the  manifesta- 
tions that  would  accompany  an  actual  experience 
of  the  evil  feared.  The  destructive  passions  are 
shown  in  a  general  tension  of  the  muscular  system, 
in  gnashing  of  the  teeth  and  protrusion  of  the 


UTILITY   OF   BODILY   CHANGES     187 

claws,  in  dilated  eyes  and  nostrils,  in  growls ;  and 
these  are  weaker  forms  of  the  actions  that  accom- 
pany the  killing  of  prey."  McDougall  ^  has  de- 
veloped this  idea  systematically  and  has  suggested 
that  an  association  has  become  established  between 
peculiar  emotions  and  peculiar  instinctive  reac- 
tions; thus  the  emotion  of  fear  is  associated  with 
the  instinct  for  flight,  and  the  emotion  of  anger 
or  rage  with  the  instinct  for  fighting  or  attack. 
Crile  ■*  likewise  in  giving  recent  expression  to 
the  same  view  has  emphasized  the  importance 
of  adaptation  and  natural  selection,  operative 
through  myriads  of  years  of  racial  experience,  in 
enabling  us  to  account  for  the  already  channeled 
responses  which  we  find  established  in  our  nervous 
organization.  And  on  a  principle  of  "i^hylogenetic 
association"  he  assumes  that  fear,  born  of  innu- 
merable injuries  in  the  course  of  evolution,  has  de- 
veloped into  portentous  foreshadowing  of  possible 
injury  and  has  become,  therefore,  capable  of  arous- 
ing in  the  bodj^  all  the  offensive  and  defensive 
activities  that  favor  the  survival  of  the  organism. 
Because  the  increase  of  adrenin  and  the  increase 
of  sugar  in  the  blood,  following  painful  or  strong 
emotional  experiences,  are  reflex  in  character,  and 
because  reflexes  as  a  rule  are  useful  responses, 
we  are  justified  in  the  assumption  that  under  these 
circumstances  these  reactions  are  useful.  What, 
then,  is  their  possible  value? 


188  BODILY    CHANGES 

In  order  that  these  reactions  may  be  useful 
they  must  be  prompt.  Such  is  the  case.  Some 
observations  made  by  one  of  my  students,  Mr.  H. 
Osgood,  show  that  the  latent  period  of  adrenal 
secretion,  when  the  splanchnic  nerve  is  stimulated 
below  the  diaphragm,  is  not  longer  than  16  sec- 
onds ;  and  Macleod  ^  states  that  within  a  few  min- 
utes after  splanchnic  stimulation  the  sugar  in  the 
blood  rises  between  10  and  30  per  cent.  The  two 
secretions  are,  therefore,  almost  instantly  ready 
for  service. 

Conceivably  the  two  secretions  might  act  in  con- 
junction, or  each  might  have  its  own  function  alone. 
Thus  adrenin  might  serve  in  cooperation  with 
nervous  excitement  to  produce  increase  of  blood 
sugar,  or  it  might  have  that  function  and  other 
functions  quite  apart  from  that.  Before  these 
possibilities  are  considered,  however,  the  value  of 
the  increased  blood  sugar  itself  will  be  discussed. 

The  Utility  of  the  Increased  Blood  Sugar  as  a  Source  of 
Muscular  Energy 

When  we  were  working  on  emotional  glycosuria 
a  clue  to  the  significance  of  the  increase  of  sugar  in 
the  blood  was  found  in  McDougall's  suggestion  of 
a  relation  between  "flight  instinct"  and  "fear 
emotion,"  and  "pugnacity  instinct"  and  "anger 
emotion."  And  the  point  was  made  that,  since 
the  fear  emotion  and  the  anger  emotion  are,  in 


UTILITY   OF   BODILY   CHANGES     189 

wild  life,  likely  to  be  followed  by  activities  (run- 
ning or  fighting)  which  require  contraction  of  great 
muscular  masses  in  supreme  and  prolonged  strug- 
gle, a  mobilization  of  sugar  in  the  blood  might  be 
of  signal  service  to  the  laboring  muscles.  Pain — 
and  fighting  is  almost  certain  to  involve  pain — 
would,  if  possible,  call  forth  even  greater  muscular 
effort.  "In  the  agony  of  pain  almost  every  muscle 
of  the  body  is  brought  into  strong  action,"  Dar- 
win ^  wrote,  for  "great  pain  urges  all  animals, 
and  has  urged  them  during  endless  generations, 
to  make  the  most  violent  and  diversified  efforts 
to  escape  from  the  cause  of  suffering."* 

•  It  is  recognized  that  both  pain  and  the  major  emotions 
may  have  at  times  depressive  rather  than  stimulating  effects. 
For  example,  Martin  and  Lacey  have  shown  (American  Jour- 
nal of  Physiology,  1914,  xxxiii,  p.  212)  that  such  stimuli  as 
would  induce  pain  may  cause  a  fall  of  blood  pressure,  and 
they  suggest  that  the  rise  of  blood  pressure  commonly  report- 
ed at  times  of  painful  experience  is  due  to  the  psychic  dis- 
turbance that  is  simultaneously  aroused.  Conceivably  there 
is  a  relation  between  recognizing  the  possibility  of  escape 
(with  the  psychic  consequences  of  that  possibility)  and  the 
degree  of  stimulating  effect.  Thus  pains  originating  from 
the  interior  of  the  body,  or  from  injuries  sure  to  be  made 
more  painful  by  action,  would  not  likely  lead  to  action.  On 
the  other  hand,  the  whip  and  spur  illustrate  the  well-known 
excitant  effect  of  painful  stimuli. 

Similarly  in  the  case  of  the  strong  emotions,  the  effect  may 
be  paralyzing  until  there  is  a  definite  deed  to  perform.  Thus 
terror  may  be  the  most  depressing  of  all  emotions,  but,  as  Dar- 
win pointed  out  (Loc.  cit.,  p.  81),  "a  man  or  animal  driven 
through  terror  to  desperation  is  endowed  with  wonderful 
strength,  and  is  notoriously  dangerous  in  the  highest  degree." 


190  BODILY    CHANGES 

That  muscular  work  is  performed  by  energy 
supplied  in  carbonaceous  material  is  sbown  by  the 
great  increase  of  carbon-dioxide  output  in  severe 
muscular  work,  which  may  exceed  twenty  times 
the  output  during  rest.  Furthermore,  the  storage 
of  glycogen  in  muscle,  and  the  disappearance  of 
this  glycogen  deposit  from  excised  muscle  stimu- 
lated to  activity,^  or  its  reduction  after  excessive 
contractions  produced  by  strychnine,^  and  the 
lessened  ability  of  muscles  to  work  if  their  glyco- 
gen store  has  been  reduced,^  and  the  simple 
chemical  relation  between  sugar  and  the  lactic  acid 
which  appears  when  muscles  are  repeatedly  made 
to  contract,  are  all  indications  that  carbohydrate 
(sugar  and  glycogen)  is  the  elective  source  of  en- 
ergy for  contraction.  This  conclusion  is  sup- 
ported in  recent  careful  studies  by  Benedict 
and  Cathcart,^^  who  have  shown  that  a  small  but 
distinct  increase  in  the  ratio  between  the  carbon- 
dioxide  breathed  out  and  the  oxygen  breathed  in 
during  a  given  period  (the  respiratory  quotient) 
occurs  during  muscular  work,  and  that  a  decrease 
in  the  quotient  follows,  thus  pointing  to  a  larger 
proportion  of  carbohydrate  burned  during  mus- 
cular work  than  before  or  after — i.  e.,  a  call  on  the 
carbohydrate  deposits  of  the  body. 

Whether  circulating  sugar  can  be  immediately 
utilized  by  active  muscles  has  been  a  subject  of  dis- 
pute.    The  claim  of  Chauveau  and  Kaufmann^^ 


UTILITY   OF   BODILY   CHANGES     191 

that  a  muscle  uses  about  three  and  a  half  times 
as  much  blood  sugar  when  active  as  when  rest- 
ing, although  supported  by  Quinquaud/-  and 
by  Morat  and  Dufourt,^^  has  been  denied  by 
Pavy/^  who  failed  to  find  any  difference  be- 
tween the  sugar  content  of  arterial  and  venous 
blood  when  the  muscle  was  contracting ;  and  also 
by  Magnus-Lev^%^^  who  has  estimated  that  the 
amount  of  change  in  sugar  content  of  the  blood 
passing  through  a  muscle  must  be  so  slight  as  to 
be  within  the  limits  of  the  error  of  analysis.  On 
the  other  hand,  when  blood  or  Ringer's  solution 
is  repeatedly  perfused  through  contracting  heart 
muscle,  the  evidence  is  clear  that  the  contained 
sugar  may  more  or  less  completely  disappear. 
Thus  Locke  and  Rosenheim  ^'^  found  that  from 
5  to  10  centigrams  of  dextrose  disappeared  from 
Ringer's  solution  repeatedly  circulated  through 
the  rabbit  heart  for  eight  or  nine  hours.  And 
recently  Patterson  and  Starling  ^^  have  shown 
that  if  blood  is  perfused  repeatedly  through  a 
heart-lung  preparation  for  three  or  four  hours, 
and  the  heart  is  continually  stimulated  by  adrenin 
added  to  the  blood,  the  sugar  in  the  blood  wholly 
vanishes ;  or  if  the  supply  of  sugar  is  maintained, 
the  consumption  may  rise  as  high  as  8  milligrams 
per  gram  of  heart  muscle  per  hour — about  four 
times  the  usual  consumption.  When  an  animal  is 
eviscerated  it  may  be  regarded  as  a  preparation 


192  BODILY   CHANGES 

in  which  the  muscles  are  perfused  with  their  proper 
blood,  pumped  by  the  heart  and  oxygenated  by 
the  lungs.  Under  these  circumstances,  the  per- 
centage of  sugar  in  the  blood  steadily  falls/* 
because  the  utilization  by  the  tissues  is  not  com-- 
pensated  for  by  further  supply  from  the  liver. 
Thus,  although  there  may  be  doubt  that  analyses 
of  sugar  in  the  blood  flowing  into  and  out  from 
an  active  muscle  during  a  brief  period  can  be  accu- 
rate enough  to  prove  a  clear  difference,  the  evi- 
dence from  the  experiments  above  cited  shows  that 
when  the  supply  of  sugar  is  limited  it  disappears 
to  a  greater  or  less  degree  when  passed  repeatedly 
through  muscular  organs. 

The  argument  may  be  advanced,  of  course,  that 
the  sugar  which  thus  disappears  is  not  directly 
utilized,  but  must  first  be  changed  to  glycogen. 
There  is  little  basis  for  this  assumption.  There 
is,  on  the  other  hand,  considerable  evidence  that 
increasing  the  blood  sugar  does,  in  fact,  directly 
increase  muscular  efficiency.  Thus  Locke  ^^  proved 
that  if  oxygenated  salt  solution  is  perfused 
through  the  isolated  rabbit  heart,  the  beats  begin 
to  weaken  after  one  or  two  hours ;  but  if  now  0.1 
per  cent  dextrose  is  added  to  the  perfusing  liquid, 
the  beats,  at  once  become  markedly  stronger  and 
may  continue  with  very  slow  lessening  of  strength 
as  long  as  seven  hours.  And  Schumberg  ^^  noted 
that  when  he  performed  a  large  amount  of  gen- 


UTILITY   OF   BODILY   CHANGES     193 

eral  bodily  work  (thus  using  up  blood  sugar) 
and  then  tested  flexion  of  the  middle  finger  in  an 
ergograph,  the  ability  of  the  muscle  was  greater 
if  he  drank  a  sugar  solution  than  if  he  drank  an 
equally  sweet  solution  of  "dulein,"  He  did  not 
know  during  the  experiment  which  solution  he  was 
drinking.  These  observations  have  been  confirmed 
by  Prantner  and  Stowasser,  and  by  Frentzel.-^ 
In  experiments  on  cats,  Lee  and  Harrold  --  found 
that  when  sugar  is  removed  from  the  animal  by 
means  of  phlorhizin  the  tibialis  anticus  is  quick- 
ly fatigued ;  but  if,  after  the  phlorhizin  treatment, 
the  animal  is  given  an  abundance  of  sugar  and  then 
submitted  to  the  test,  the  muscle  shows  a  much 
larger  capacity  for  work.  All  this  evidence  is, 
of  course,  favorable  to  the  view  that  circulating 
sugar  may  be  quickly  utilized  by  contracting 
muscles. 

From  the  experimental  results  presented  above 
it  is  clear  that  muscles  work  preferably  by  utilizing 
the  energy  stored  in  sugar,  that  great  muscular 
labor  is  capable  of  considerably  reducing  the  quan- 
tity of  stored  glycogen  and  of  circulating  sugar, 
and  that  under  circumstances  of  a  lessened  sugar 
content  the  increase  of  blood  sugar  considerably 
augments  the  ability  of  muscles  to  continue  con- 
tracting. The  conclusion  seems  justified,  there- 
fore, that  the  increased  blood  sugar  attendant  on 
the  major  emotions  and  pain  would  be  of  direct 


194  BODILY   CHANGES 

benefit  to  the  organism  in  the  strenuous  muscular 
efforts  involved  in  flight  or  conflict  or  struggle  to 
be  free. 

The  Utility  of  Increased  Adrenin  in  the  Blood  as  an 
Antidote  to  the  Effects  of  Fatigue 

The  function  which  the  discharged  adrenin  itself 
might  have  in  favoring  vigorous  muscular  con- 
traction has  already  been  suggested  in  the  chapter 
on  the  effect  of  adrenin  in  restoring  the  irritability 
of  fatigued  muscle.  Some  of  the  earliest  evidence 
proved  that  removal  of  the  adrenal  glands  has  a 
debilitating  effect  on  muscular  power,  and  that 
injection  of  adrenal  extract  has  an  invigorating 
effect.  For  these  reasons  it  seemed  possible  that 
increased  adrenal  secretion,  as  a  reflex  result  of 
pain  or  the  major  emotions,  might  act  in  itself  as 
a  dynamogenic  factor  in  the  performance  of  mus- 
cular work.  It  was  on  the  basis  of  that  possibility 
that  Nice  and  I  tested  the  effect  of  stimulating 
the  splanchnic  nerves  (thus  causing  adrenal  secre- 
tion), or  injecting  adrenin,  on  the  contraction  of 
the  fatigued  tibialis  anticus.  We  found,  as  already 
described,  that  when  arterial  pressure  was  of  nor- 
mal height,  and  was  prevented  from  rising  in  the 
legs  while  the  splanchnic  was  being  stimulated, 
there  was  a  distinct  rise  in  the  height  of  contrac- 
tion of  the  fatigued  muscle.  And  we  drew  the 
Inference  that  adrenin  set  free  in  the  blood  may 


UTILITY   OF   BODILY   CHANGES     195 

operate  favorably  to  the  organism  by  preparing 
fatigued  muscles  for  better  response  to  the  nervous 
discharges  sent  forth  in  great  excitement. 

This  inference  led  to  the  experiments  by  Gruber, 
who  examined  the  effects  of  minute  amounts  of 
adrenin  (0.1  or  0.5  cubic  centimeter,  1:100,000), 
and  also  of  splanchnic  stimulation,  on  the  thresh- 
old stimulus  of  fatigued  neuro-muscular  and  mus- 
cular apparatus.  Fatigue,  the  reader  will  recall, 
raises  the  threshold  not  uncommonly  100  or  200 
per  cent,  and  in  some  instances  as  much  as  600  per 
cent.  Rest  will  restore  the  normal  threshold  in 
periods  varying  from  fifteen  minutes  to  two  hours, 
according  to  the  length  of  previous  stimulation. 
If  a  small  dose  of  adrenin  is  given,  however,  the 
normal  threshold  may  be  restored  in  three  to  five 
minutes. 

From  the  foregoing  evidence  the  conclusion  is 
warranted  that  adrenin,  when  freely  liberated  in 
the  blood,  not  only  aids  in  bringing  out  sugar  from 
the  liver's  store  of  glycogen,  but  also  has  a  remark- 
able influence  in  quickly  restoring  to  fatigued  mus- 
cles, which  have  lost  their  original  irritability,  the 
same  readiness  for  response  which  they  had  when 
fresh.  Thus  the  adrenin  set  free  in  pain  and  in 
fear  and  rage  would  put  the  muscles  of  the  body 
unqualifiedly  at  the  disposal  of  the  nervous  sys- 
tem; the  difficulty  which  nerve  impulses  might 
have  in  calling  the  muscles  into  full  activity  would 


196  BODILY   CHANGES 

be  practically  abolished ;  and  this  provision,  along 
with  the  abundance  of  energy-supplying  sugar 
newly  flushed  into  the  circulation,  would  give  to 
the  animal  in  which  these  mechanisms  are  most 
efficient  the  best  possible  conditions  for  putting 
forth  supreme  muscular  efforts.* 

The  Question  Whether  Adrenin  Normally  Secreted 
Inhibits  the  Use  of  Sugar  in  the  Body 

The  only  evidence  opposed  to  the  conclusion 
which  has  just  been  drawn  is  that  which  may  be 
found  in  results  recently  reported  by  Wilenko.  He 
injected  adrenin  into  urethanized  rabbits,  usually 
one  milligram  per  kilo  body  weight,  and  then  found 
that  the  animals  did  not  oxidize  any  part  of  an 
intravenous  injection  of  glucose.  Rabbits  supplied 
with  glucose  in  a  similar  manner,  but  not  given 
adrenin,  have  an  increased  respiratory  quotient. 
Wilenko  -^  concluded,  therefore,  that  adrenin  les- 
sens the  capacity  of  the  organism  to  burn  carbo- 
hydrates. In  a  later  paper  he  reported  that  adren- 
in, when  added,  with  glucose,  to  physiological  salt 
solution  (Locke's),  and  perfused  through  the  iso- 
lated rabbit  heart,  notably  increases  the  use  of 
sugar  by  the  heart  (from  2.2-2.8  to  2.9-4.3  milli- 

*  If  these  results  of  emotion  and  pain  are  not  "worked  off" 
by  action,  it  is  conceivable  that  the  excessive  adrenin  and 
sugar  in  the  blood  may  have  pathological  effects.  (Cf.  Can- 
non :  Journal  of  the  American  Medical  Association,  1911,  Ivi, 
p.  742.) 


UTILITY   OF   BODILY   CHANGES     197 

grams  of  glucose  per  gram  of  heart  muscle  per 
hour),  but  that  the  heart  removed  after  the  animal 
has  received  a  subcutaneous  injection  of  adrenin 
uses  much  less  sugar,  only  0.5-1.2  milligrams 
per  gram  per  hour.  From  these  results  Wilen- 
ko  -^  concludes  that  the  glycosuria  following  in- 
jection of  adrenin  is  the  result  of  disturbance  of 
the  use  of  sugar — an  effect  which  is  not  direct  on 
the  sugar-consuming  organ,  but  indirect  through 
action  on  some  other  organ. 

Wilenko's  conclusion  fails  to  account  readily  for 
the  disappearance  of  glycogen  from  the  liver  in 
adrenin  glycosuria.  Furthermore,  Lusk  -^  has 
recently  reported  that  the  subcutaneous  adminis- 
tration of  adrenin  (one  milligram  per  kilo  body 
weight)  to  dogs,  simultaneously  with  50  grams  of 
glucose  by  mouth,  interferes  not  at  all  with  the 
use  of  the  sugar — the  respiratory  quotient  remains 
for  several  hours  at  1.0;  i.e.,  at  the  figure  which 
glucose  alone  would  have  given.  In  other  words, 
Lusk's  results  with  dogs  are  directly  contradictory 
to  Wilenko's  results  with  rabbits.  Nevertheless, 
Wilenko's  conclusion  might  be  quite  true  for  the 
glycosuria  produced  by  adrenin  alone  (which  must 
be  excessive),  and  yet  have  no  bearing  whatever 
on  the  glycosuria  produced  physiologically  by 
splanchnic  stimulation,  even  though  some  adrenin 
is  thereby  simultaneously  liberated. 

The  amount  of  injected  adrenin  used  to  produce 


198  BODILY    CHANGES 

adrenin  glycosuria  is  enormous.  Osgood  has  stud- 
ied in  the  Harvard  Physiological  Laboratory  the 
effects  on  blood  pressure  of  alternately  stimulating 
the  left  splanchnic  nerves  (with  the  splanchnic 
vessels  eliminated)  and  injecting  adrenin,  and  by 
this  method  of  comparison  ^^  has  shown  that 
the  amount  secreted  after  five  seconds  of  stimula- 
tion varies  betweet  0.0015  and  0.007  milligram.  If 
0.005  milligram  is  taken  as  a  rather  high  average 
figure,  and  doubled  (for  two  glands),  the  amount 
would  be  0.01  milligram.  To  produce  adrenin  gly- 
cosuria, an  animal  weighing  two  kilos  would  be 
injected  with  two  hundred  times  this  amount.  It 
is  granted  that  more  adrenin  would  be  secreted  if 
the  nerves  were  stimulated  longer  than  five  sec- 
onds, and  that  with  injection  under  the  skin  or  into 
the  abdominal  cavity  (to  produce  glycosuria),  the' 
amount  of  adrenin  in  the  blood  at  one  time  would 
not  be  so  great  as  if  the  injection  were  into  a  vein ; 
but  even  with  these  concessions  the  amount  of  ad- 
renin in  the  blood,  when  it  has  been  injected  to 
produce  glycosuria,  is  probably  very  much  above 
the  amount  following  physiological  stimulation  of 
the  glands. 

Other  evidence  that  the  amount  of  adrenin  dis- 
charged when  the  glands  are  stimulated  is  not  so 
great  as  the  amount  needed  to  produce  glycosuria 
when  acting  alone  is  presented  in  experiments  by 
Macleod.2^     He  found   that  if  the  nerve  fibres 


UTILITY   OF   BODILY   CHANGES     199 

to  the  liver  were  destroyed,  stimulation  of  the 
splanchnic,  which  would  cause  increased  adrenal 
secretion,  did  not  increase  the  blood  sugar.  The 
increased  blood  sugar  due  to  splanchnic  stimula- 
tion, therefore,  is  a  nervous  effect,  dependent,  to 
be  sure,  on  the  presence  of  adrenin  in  the  blood, 
but  the  amount  of  adrenin  present  is  not  in  itself 
capable  of  evoking  increase. 

■  Furthermore,  the  increased  blood  sugar  follow- 
ing splanchnic  stimulation  may  long  outlast  the 
stimulation  period.  The  adrenals,  however,  as  has 
been  demonstrated  by  Osgood,  are  soon  fatigued, 
and  fail  to  respond  to  repeated  stimulation.  They 
seem  to  be  incapable  of  prolonged  action. 

Again,  as  Macleod  -^  has  shown,  a  rise  in  the 
sugar  content  of  the  blood  can  be  induced,  if  the 
adrenals  are  intact,  merely  by  stimulating  the 
nerves  going  to  the  liver.  The  increased  blood 
sugar  of  splanchnic  origin,  therefore,  is  not  due  to 
a  disturbance  of  the  use  of  sugar  in  the  body,  as 
Wilenko  claims  for  the  increase  after  adrenin  in- 
jection, but  is  a  result  of  a  breaking  down  of  the 
stored  glycogen  in  the  liver  and  is  of  nervous 
origin. 

We  may  conclude,  therefore,  that  since  the  condi- 
tions of  Wilenko's  observations  are  not  compar- 
able with  emotional  conditions,  his  inferences  are 
not  pertinent  to  the  present  discussion;  that  when 
both  adrenin  and  sugar  are  increased  in  the  blood 


200  BODILY   CHANGES 

as  a  result  of  excitement,  the  higher  percentage 
of  sugar  is  not  due  to  adrenin  inhibiting  the  use 
of  sugar  by  the  tissues,  and  that  there  is  no  evi- 
dence at  present  to  show  that  the  brief  augmenta- 
tion of  adrenal  discharge,  following  excitement  or 
splanchnic  stimulation,  affects  in  any  deleterious 
manner  the  utilization  of  sugar  as  a  source  of  en- 
ergy. Indeed,  the  observation  of  Wilenko  and  of 
Patterson  and  Starling,  above  mentioned,  that  ad- 
renin increases  the  use  of  sugar  by  the  heart,  may 
signify  that  a  physiological  discharge  of  the  ad- 
renals can  have  a  favorable  rather  than  an  unfa- 
vorable effect  on  the  employment  of  sugar  by  the 
tissues. 

The  Vascular  Changes  Produced  by  Adrenin  Favorable  to 
Supreme  Muscular  Exertion 

Quite  in  harmony  with  the  foregoing  argument 
that  sugar  and  adrenin,  which  are  poured  into  the 
blood  during  emotional  excitement,  render  the  or- 
ganism more  efficient  in  the  physical  struggle  for 
existence,  are  the  vascular  changes  wrought  by 
increased  adrenin,  probably  in  cooperation  with 
sympathetic  innervations.  The  studies  of  volume  x^ 
changes  of  parts  of  the  body,  made  by  Oliver  and 
Schafer,  have  already  been  mentioned.  Their  ob- 
servations, it  will  be  remembered,  showed  that 
injected  adrenin  drove  the  blood  from  the  abdom- 
inal viscera  into  the  organs  called  upon  in  emer- 


UTILITY   OF   BODILY   CHANGES     201 

gencies — into  the  central  nervous  system,  the 
lungs,  the  heart,  and  the  active  skeletal  muscles. 
The  absence  of  effective  vasoconstrictor  nerves 
in  the  brain  and  the  lungs,  and  the  dilation  of 
vessels  in  the  heart  and  skeletal  muscles  during 
times  of  increased  activity,  make  the  blood  supply 
to  these  parts  dependent  on  the  height  of  general 
arterial  pressure.  In  pain  and  great  excitement, 
as  we  have  already  noted,  this  pressure  is  likely 
to  be  much  elevated,  and  consequently  the  blood 
flow  through  the  unconstricted  or  actually  dilated 
vessels  of  the  body  will  be  all  the  more  abundant. 
Adrenin  has  a  well-known  stimulating  effect  on 
the  isolated  heart — causing  an  increase  both  in  the 
rate  and  the  amplitude  of  cardiac  contraction.  This 
effect  accords  with  the  general  rule  that  adrenin 
simulates  the  action  of  sympathetic  impulses.  It 
is  commonly  stated,  however,  that  if  the  heart 
holds  its  normal  relations  in  the  body,  adrenin 
causes  slowing  of  the  beat.-^  This  view  is  doubt- 
less due  to  the  massive  doses  that  have  been 
employed,  which  are  quite  beyond  physiological 
limits  and  which  induce  such  enormous  increases 
of  arterial  pressure  that  the  natural  influence  of 
adrenin  on  heart  muscle  is  overcome  by  mechanical 
obstacles  to  quick  contractions  and  by  inhibitory 
impulses  from  the  central  nervous  system.  IIos- 
kins  and  Lovellette  have  recently  shown  that  when 
the  precaution  is  taken  to  inject  adrenin  into  a  vein 


202  BODILY   CHANGES 

in  a  manner  resembling  the  discharge  from  the 
adrenal  glands,  not  only  is  there  increased  blood 
pressure,  bnt  generally,  also,  an  acceleration  of  the 
pulse.^^  At  the  same  time,  therefore,  that  a 
greater  amount  of  work,  from  increased  arterial 
pressure,  is  demanded  of  the  heart,  blood  is  de- 
livered to  the  heart  in  greater  abundance,  and  the 
muscle  is  excited  to  more  rapid  and  vigorous  pul- 
sations. The  augmentation  of  the  heart  beat  is 
thus  coordinate  with  the  other  adaptive  functions 
of  the  adrenal  glands  in  great  emergencies. 

The  Changes  in  Respiratory  Functon  Also  Favorable 
TO  Great  Effort 

The  urgent  need  in  struggle  or  flight  is  a  gen- 
erous supply  of  oxygen  to  oxidize  the  metabolites 
of  muscular  contraction,  and  a  quick  riddance  of 
the  resultant  carbon-dioxide  from  the  body.  The 
moment  vigorous  exercise  is  begun  the  breathing 
at  once  changes  so  as  to  bring  about  a  more  thor- 
ough ventilation  of  the  lungs.  And  one  of  the  most 
characteristic  reactions  of  animals  in  pain  and 
emotional  excitement  is  deep  and  rapid  respiration. 
Again  the  reflex  response  is  precisely  what  would 
be  most  serviceable  to  the  organism  in  the  stren- 
uous efforts  of  fighting  or  escape  that  might  accom- 
pany or  follow  distress  or  fear  or  rage.  It  is 
known  that  by  such  forced  respirations  the  carbon- 
dioxide  content  of  the  blood  can  be  so  much  re- 


UTILITY   OF   BODILY   CHANGES     203 

duced  that  the  need  for  any  breathing  whatever 
may  be  deferred  for  as  much  as  a  minute  or 
even  longer.^^  And  Douglas  and  Haldane  ^-  have 
found  that  moderately  forced  breathing  for  three 
minutes  previous  to  severe  muscular  exertion  re- 
sults in  greatly  diminishing  the  subsequent  res- 
piratory distress,  as  well  as  lessening  the  amount 
of  air  breathed  and  the  amount  of  carbon-dioxide 
given  off.  Furthermore,  the  heart  beats  less  rap- 
idly after  the  performance  and  returns  more 
quickly  from  its  increased  rate  to  normal.  The 
forced  respirations  in  deeply  emotional  experi- 
ences can  be  interpreted,  therefore,  as  an  antici- 
patory reduction  of  the  carbon-dioxide  in  the  blood, 
a  preparation  for  the  augmented  discharge  of 
carbon-dioxide  into  the  blood  as  soon  as  great 
muscular  exertion  begins.* 

As  the  air  moves  to  and  fro  in  the  lungs  with 
each  respiration,  it  must  pass  through  the  fine 
divisions  of  the  air  tubes  or  bronchioles.  The 
bronchioles  are  provided  with  smooth  muscle, 
which,  in  all  probability,  like  smooth  muscle  else- 
where in  the  body,  is  normally  held  in  a  state  of 

•  The  excessive  production  of  heat  in  muscular  work  gives 
rise  to  sweating.  The  evaporation  of  sweat  helps  to  keep  the 
body  temperature  from  rising  unduly  from  the  heat  of  exer- 
tion. Again  in  strong  emotion  and  in  pain  the  "cold  sweat" 
that  appears  on  the  skin  may  be  regarded  as  a  reaction 
anticipatory  of  the  strenuous  muscular  movements  that  are 
likely  to  ensue. 


204  BODILY   CHANGES 

tonic  contraction.  When  this  tonic  contraction  is 
much  increased,  as  in  asthma,  breathing  becomes 
difficult,  and  even  with  the  body  at  rest  unusual 
effort  is  then  required  to  maintain  the  minimaP 
necessary  ventilation  of  the  lungs.  During  stren- 
uous exertion,  with  each  breath  the  air  must  rush 
through  the  bronchioles  in  greatly  increased  vol- 
ume and  speed.  Thus  in  a  well  person  "winded" 
with  running,  for  example,  the  bronchioles  might 
become  relatively  too  small  for  the  stream  of  air, 
just  as  they  are  too  small  in  a  person  ill  with 
asthma.  And  then  some  extra  energy  would  have 
to  be  expended  to  force  the  air  back  and  forth  with 
sufficient  rapidity  to  satisfy  the  bodily  needs.  It 
is  probable  that  even  under  the  most  favorable  con- 
ditions, the  labored  breathing  in  hard  exercise  in- 
volves to  some  degree  the  work  of  accelerating  the 
tidal  flow  of  the  respiratory  gases.  This  extra 
labor  would  obviously  be  reduced,  if  the  tonic  con- 
traction of  the  ring-muscles  in  the  wall  of  the 
bronchioles  was  reduced,  so  that  the  tubules  were 
enlarged.  It  has  been  shown  by  a  number  of  in- 
vestigators, who  have  used  various  methods,  that 
adrenin  injected  into  the  blood  stream  has  as  one 
of  its  precise  actions  the  dilating  of  the  bronchi- 
oles.^^ The  adrenin  discharged  in  emotional  ex- 
citement goes  to  the  lungs  before  entering  into 
relation  with  any  other  organ  except  the  right 
heart  chamber;  it  may,  therefore,  have  as  its  first 


UTILITY   OF   BODILY   CHANGES     205 

effect  the  relaxation  of  the  smooth  muscles  of  the 
lungs.  This  would  be  another  very  direct  means 
of  rendering  the  organism  more  efficient  when 
tierce  struggle  calls  for  a  bounteous  supply  of  fresh 
air  and  a  speedy  discharge  of  the  carbonaceous 
waste. 


Effects  Produckd  in  Asphyxia  Similar  to  those  Produced 
IN  Pain  and  Excitement 

All  the  bodily  responses  occurring  in  pain  and 
emotional  excitement  have  thus  far  been  consid- 
ered as  anticipatory  of  the  instinctive  acts  which 
naturally  follow.  And  as  we  have  seen,  these  re- 
sponses can  reasonably  be  interpreted  as  prepar- 
atory to  the  great  exertions  which  may  be  de- 
manded of  the  organism.  This  interpretation  of 
the  facts  is  supported  by  the  discovery  that  a 
mechanism  exists  whereby  the  changes  initiated 
in  an  anticipatory  manner  by  emotional  excite- 
ment are  continued  or  perhaps  augumented  by  the 
exertion  itself. 

Great  exertion,  such  as  might  attend  flight  or  yr 
conflict,  would  result  in  an  excessive  production  of  ' 
carbon-dioxide.  Then,  although  respiratory  and 
circulatory  changes  of  emotional  origin  may  have 
prepared  the  body  for  struggle,  the  emotional  pro- 
visions for  keeping  the  working  parts  at  a  high 
level  of  efficiency  may  not  continue  to  operate,  or 


206  BODILY    CHANGES 

they  may  not  be  adequate.  If  there  is  painful  gasp- 
ing for  breath  in  the  course  of  prolonged  and  vig- 
orous exertion,  or  for  a  considerable  period  after 
the  work  has  ceased,  a  condition  of  partial 
asphyxia  has  evidently  been  induced.  This  condi- 
tion, as  everyone  knows,  is  distinctly  unfavorable 
to  further  effort.  But  the  asphyxia  itself  may  act 
as  a  stimulus.^^ 

In  our  examination  of  the  influence  of  various 
conditions  on  the  secretion  of  the  adrenal  glands, 
Hoskins  and  I^^  tested  the  effects  of  asphyxia. 
By  use  of  the  intestinal  segment  as  an  indicator  we 
compared  the  action  of  blood,  taken  as  nearly  si- 
multaneously as  possible  from  the  vena  cava  above 
the  adrenal  vessels  and  from  the  femoral  vein  be- 
fore asphyxia,  with  blood  taken  from  the  same 
sources  after  asphyxia  had  been  produced.  The 
femoral  venous  blood  after  passing  the  capillaries 
of  the  leg  thus  acted  as  a  standard  for  the  same 
blood  after  receiving  the  contribution  of  the 
adrenal  veins.  Asphyxia  was  caused  by  covering 
the  tracheal  cannula  until  respiration  became 
labored  and  slow,  but  capable  of  recovery  when  air 
was  admitted.  It  may  be  regarded,  therefore,  as 
not  extreme. 

The  results  of  the  degree  of  asphyxia  above 
described  are  shown  by  graphic  record  in  Fig.  36. 
Blood  taken  from  the  vena  cava  and  from  the 
femoral  vein  before  asphyxia  ("normal")  failed  to 


UTILITY   OF   BODILY   CHANGES     207 

cause  inhibition  of  the  contractions.  Blood  taken 
from  the  femoral  vein  after  asphyxia  produced  al- 
most the  same  effect  as  blood  from  the  same  vein 
before;  asphyxia,  therefore,  had  wrought  no 
change  demonstrable  in  the  general  venous  flow. 


Figure  36. — Adrenal  secretion  produced  by  as- 
phyxia. At  1  normal  vena-cava  blood  applied, 
at  2  removed.  At  3  normal  blood  from  fomoral 
vein  applied,  at  4  removed.  At  5  blood  from 
femoral  vein  after  asphyxia  applied,  at  6  re- 
moved. At  7  blood  from  the  vena  cava  after 
asphyxia  applied.     Time,  half-minutes. 


Blood  taken  from  the  vena  cava  after  asphyxia 
had,  on  the  contrary,  an  effect  markedly  unlike 
blood  from  the  same  region  before  (compare  the 
record  after  1  and  after  7,  Fig.  36) — it  caused  the 


208  BODILY   CHANGES 

typical  inhibition  whicli  indicates  the  presence  of 
adrenal  secretion.* 

That  the  positive  result  obtained  in  moderate 
asphyxia  is  not  attributable  to  other  agencies  in 
the  blood  than  adrenin  is  indicated  by  the  failure 
of  asphyxial  femoral  blood  to  cause  inhibition, 
while  vena-cava  blood,  taken  almost  simultane- 
ously, brought  about  immediate  relaxation  of  the 
muscle.  The  conclusion  was  drawn,  therefore,  ~ 
that  asphyxia  results  in  increased  secretion  of  the 
adrenal  glands. 

This  conclusion  has  been  supported  by  Bor- 
berg  and  Fridericia,^^  and  also  by  Starkenstein,^'^ 
who  found  that  an  increase  of  carbon-dioxide  in 
the  blood  lessens  the  adrenin  in  the  adrenal  me- 

*  This  positive  result  might  suggest  that  the  comparison  of 
both  femoral  and  vena-cava  blood  under  each  condition  was 
unnecessary,  and  that  a  comparison  merely  of  vena-cava 
blood  before  and  after  asphyxia  would  be  sufficient.  Positive 
results  were  indeed  thus  secured,  but  they  occurred  even 
when  the  adrenal  glands  were  carefully  removed  and  extreme 
asphyxia  (i.  e.,  stoppage  of  respiration)  was  induced.  That 
the  blood  may  contain  in  extreme  asphyxia  a  substance  or 
substances  capable  of  causing  inhibition  of  intestinal  con- 
tractions was  thus  demonstrated.  In  one  instance,  after  the 
blood  was  proved  free  from  adrenin,  the  aorta  and  vena  cava 
were  tied  close  below  the  diaphragm,  and  the  carotids  were 
tied  about  midway  in  the  neck.  Extreme  asphyxia  was 
produced  (lasting  five  minutes).  Blood  now  taken  from  the 
heart  caused  marked  inhibition  of  the  beating  intestinal 
segment.  Probably,  therefore,  the  inhibitory  action  of  blood 
taken  from  an  animal  when  extremely  asphyxiated  cannot  be 
due  to  adrenin  alone. 


UTILITY   OF   BODILY    CHANGES     209 

diilla.  And  recently  Czubalski  ^^  also  has  inferred, 
from  the  rise  of  blood  pressure  in  asphyxia  when 
the  adrenals  are  Latact  and  the  absence  of  the  rise 
if  the  adrenals  are  removed,  that  asphyxia  sets 
free  adrenin  in  the  blood. 

Asphyxia,  like  pain  and  excitement,  not  only  lib- 
erates adrenin,  but,  as  might  be  inferred  from  that 
fact,  also  mobilizes  sugar.^*^  And,  furthermore, 
Starkenstein  ^^  has  shown  that  the  asphyxia  due 
to  carbon-monoxide  poisoning  is  not  accompanied 
by  increased  blood  sugar  if  the  adrenal  glands  have 
been  removed. 

In  case  strong  emotions  are  followed  by  vigor- 
ous exertions,  therefore,  asphyxia  is  likely  to 
result,  and  this  will  act  in  conjunction  with  the 
emotional  excitement  and  pain,  or  perhaps  in  con- 
tinuation of  the  influences  of  these  states,  to  bring 
forth  still  more  adrenal  discharge  and  still  further 
output  of  sugar  from  the  liver.  And  these  in  turn 
would  serve  the  laboring  muscles  in  the  manner 
already  described.  This  suggestion  is  in  accord 
with  Macleod's  *^  that  the  increased  freeing  of 
glycogen  from  the  liver  produced  by  muscular  ex- 
ercise is  possibly  associated  with  increased  carbon- 
dioxide  in  the  blood.  And  it  also  harmonizes  with 
Zuntz's  statement  ^-  that  the  asphyxia  of  great 
physical  exertion  may  call  out  sugar  to  such  a  de- 
gree that,  in  spite  of  the  increased  use  of  it  in  the 
active  muscles,  glycosuria  may  ensue. 


210  BODILY   CHANGES 

The  evidence  previously  adduced  that  adrenin 
causes  relaxation  of  the  smooth  muscle  of  the 
bronchioles,  taken  in  conjunction  with  the  evidence 
that  adrenal  secretion  is  liberated  in  asphyxia,  sug- 
gests that  relief  from  difficult  breathing  may  thus 
be  automatically  provided  for  in  the  organism. 
The  well-known  phenomenon  of  "second  wind"  is 
characterized  by  an  almost  miraculous  refreshment 
and  renewal  of  vigor,  after  an  individual  has  per- 
sisted in  violent  exertion  in  spite  of  being  "out  of 
breath."  It  seems  not  improbable  that  this  phe- 
nomenon, for  which  many  explanations  have  been 
offered,  is  really  due  to  setting  in  operation  the 
supporting  mechanism  which,  as  we  have  seen, 
plays  so  important  a  role  in  augmenting  bodily 
vigor  in  emotional  excitement.  The  release  of 
sugar  and  adrenin,  the  abundance  of  blood  flow 
through  the  muscles — supplying  energy  and  les- 
sening fatigue — and  the  relaxation  of  the  bronchi- 
olar  walls,  are  all  occurrences  which  may  reason- 
ably be  regarded  as  resulting  from  asphyxia.  And 
when  they  take  place  they  doubtless  do  much  to 
abolish  the  distress  itself  by  which  they  were  occa- 
sioned. According  to  this  explanation  "second 
wind"  would  consist  in  the  establishment  of  the 
same  group  of  bodily  changes,  leading  to  more 
efficient  physical  struggle,  that  are  observed  in 
pain  and  excitement. 


UTILITY   OF   BODILY   CHANGES     211 

The  Utility  of  Rapid  Coagulation  in  Preventenq 
Loss  OF  Blood 

The  increase  of  blood  sugar,  the  secretion  of 
adrenin,  and  the  altered  circulation  in  pain  and 
emotional  excitement  have  been  interpreted  in  the 
foregoing  discussion  as  biological  adaptations  to 
conditions  in  wild  life  which  are  likely  to  involve 
pain  and  emotional  excitement,  i.  e,,  the  necessities 
of  fighting  or  flight.  The  more  rapid  clotting  of 
blood  under  these  same  circumstances  may  also  be 
regarded  as  an  adaptive  process,  useful  to  the  or- 
ganism. The  importance  of  conserving  the  blood, 
especially  in  the  struggles  of  mortal  combat,  needs 
no  argument.  The  effect  of  local  injury  in  favor- 
ing the  formation  of  a  clot  to  seal  the  opened  ves 
sels  is  obviously  adaptive  in  protecting  the  organ- 
ism against  hemorrhage.  The  injury  that  causes 
opening  of  blood  vessels,  however,  is,  if  extensive, 
likely  also  to  produce  pain.  And,  as  already 
shown,  conditions  producing  pain  increase  adrenal 
secretion  and  hasten  coagulation.  Thus  injury 
would  be  made  less  dangerous  as  an  occasion  for 
serious  hemorrhage  by  two  effects  which  the  in- 
jury itself  produces  in  the  body — the  local  effect 
on  clotting  at  the  region  of  injury  and  the  general 
effect  on  the  speed  of  clotting  wrought  by  reflex 
secretion  of  adrenin. 

According  to  the  argument  here  presented  the 
strong  emotions,  as  fear  and  anger,  are  rightly 


212  BODILY   CHANGES 

interpreted  as  the  concomitants  of  bodily  changes 
which  may  be  of  utmost  service  in  subsequent  ac- 
tion. These  bodily  changes  are  so  much  like  those 
which  occur  in  pain  and  fierce  struggle  that,  as 
early  writers  on  evolution  suggested,  the  emotions 
may  be  considered  as  foreshadowing  the  suffering 
and  intensity  of  actual  strife.  On  this  general 
basis,  therefore,  the  bodily  alterations  attending 
violent  emotional  states  would,  as  organic  prepara- 
tions for  fighting  and  possible  injury,  naturally 
involve  the  effects  which  pain  itself  would  pro- 
duce. And  increased  blood  sugar,  increased 
adrenin,  an  adapted  circulation  and  rapid  clotting 
would  all  be  favorable  to  the  preservation  of  the 
organism  that  could  best  produce  them. 

KEFEEENCES 

^  See  Darwin :  Expression  of  Emotions  in  Man  and  Ani- 
mals, New  York,  1905,  pp.  101,  117. 

2  Spencer :  Principles  of  Psychology,  London,  1855. 

^  McDougall :  Introduction  to  Social  Psychology,  London, 
1908,  pp.  49,  59. 

*  Crile :  Boston  Medical  and  Surgical  Journal,  1910, 
clxiii,  p.  893. 

^  Macleod :  Diabetes,  etc.,  p.  80. 

^  Darwin :  Loc.  cit.,  p.  72. 

^  Nasse :  Archiv  f  iir  die  gesammte  Physiologic,  1869,  ii,  p. 
106;  1877,  xiv,  p.  483. 

^Frentzel:  Archiv  fiir  die  gesammte  Physiologie,  1894, 
Ivi,  p.  280. 

''Zuntz:  Oppenheimer's  Handbuch  der  Biochemie,  Jena, 
1911,  iv  (first  half),  p.  841. 


UTILITY    OF   BODILY    CHANGES     213 

^°  Benedict  and  Cathcart :  Muscular  Work,  a  Metabolic 
Study,  Washington,  1913,  pp.  85-87. 

^^  Chauveau  and  Kaufmann :  Comptes  Rendus,  Academie 
des  Sciences,  1886,  ciii,  p.  1062. 

12  Quinquaud :  Comptes  Rendus,  Societe  de  Biologic,  1886, 
xxxviii,  p.  410. 

13  Morat  and  Duf  ourt :  Archives  de  Physiologic,  1892, 
xxiv,  p.  327. 

i*Pavy:  The  Physiology  of  the  Carbohydrates,  London, 
1894.  p.  166. 

1^  Magnus-Levy :  v.  Noorden's  Handbueh  der  Pathologic 
des  Stoffwechsels,  1906,  i,  p.  385. 

1®  Locke  and  Rosenheim :  Journal  of  Physiology,  1907, 
xxxvi,  p.  211. 

1^ Patterson  and  Starling:  Journal  of  Physiology,  1913, 
xlvii,  p.  143. 

1*  See  Macleod  and  Pearce :  American  Journal  of  Physi- 
ology, 1913,  xxxii,  p.  192.  Pavy  and  Siau :  Journal  of  Physi- 
ology, 1903,  xxix,  p.  375.  Macleod:  American  Journal  of 
Physiology,  1909,  xxiii,  p.  278. 

1^  Locke :  Centralblatt  fiir  Physiologic,  1900,  xiv,  p.  671. 

2°  Schumberg :  Archiv  fiir  Physiologie,  1896,  p.  537. 

^1  Frentzel :  Archiv  fiir  Physiologie,  1899,  Supplement 
Band,  p.  145. 

22  Lee  and  Harrold :  American  Journal  of  Physiology, 
1900,  iv,  p.  ix. 

23  Wilenko :  Biochemische  Zeitschrif  t,  1912,  xlii,  p.  58. 

2*  Wilenko :  Archiv  fiir  experimentelle  Pathologic  und 
Pharmakologie,  1913,  Ixxi,  p.  266. 

25  Lusk :  Proceedings  of  the  Society  for  Experimental 
Biology  and  Medicine,  1914,  xi,  p.  49.  Also  Lusk  and  Riche: 
Archives  of  Internal  Medicine,  1914,  xiii,  p.  68. 

2«  See  Elliott :  Journal  of  Physiology,  1912,  xliv,  p.  376. 

2^  Macleod :  Diabetes,  etc.,  pp.  64-73. 

28  Macleod:  Diabetes,  etc.,  pp.  68-72. 

2»  See  Biedl:  Die  Innere  Sekretion,  1913,  i.  p.  464. 

^''Hoskins  and  Lovellettc:  Journal  of  the  American  Med- 
ical Association,  1914,  Lxiii,  p.  317. 


214  BODILY    CHANGES 

31  See  Haldane  and  Priestley :  Journal  of  Physiology, 
1905,  xxxii,  p.  255. 

32  Douglas  and  Haldane:  Journal  of  Physiology,  1909, 
xxxix,  p.  1. 

33  See  Januschke  and  Pollak :  Archiv  fiir  experimentelle 
Pathologie  und  Pharmakologie,  1911,  Ixvi,  p.  205.  Trendelen- 
burg :  Zentralblatt  fiir  Physiologie,  1912,  xxvi,  p.  1.  Jackson : 
Journal  of  Pharmacology  and  Experimental  Therapeutics, 
1912,  iv,  p.  59. 

3*  Cf .  Hoskins  and  McClure :  Archives  of  Internal  Medi- 
cine, 1912,  X,  p.  355. 

35  Cannon  and  Hoskins :  American  Journal  of  Physiology, 
1911,  xxix,  p.  275. 

36  Borberg :  Skandinavisches  Archiv  fiir  Physiologie,  1913, 
xxviii,  p.  125. 

3^  Starkenstein :  Zeitschrif t  fiir  experimentelle  Pathol- 
ogie und  Therapie,  1911,  x,  p.  95. 

3s  Czubalski :  Zentralblatt  fiir  Physiologie,  1913,  xxvii, 
p.  580. 

3^  Por  evidence  and  for  references  to  this  literature,  see 
Bang:  Der  Blutzucker,  Wiesbaden,  1913,  pp.  104-108. 

^°  Starkenstein :  Loc.  cib.,  p.  94. 

*i  Macleod :  Diabetes,  etc.,  p.  184. 

^2  Zuntz :  Loc.  cit.,  p.  854. 


-     CHAPTER    XII 

THE    ENEEGIZING    INFLUENCE    OF    EMOTIONAL 
EXCITEMENT 

The  close  relation  between  emotion  and  muscu- 
lar action  has  long  been  perceived.  As  Sher- 
rington ^  has  pointed  out,  "Emotion  'moves'  us, 
hence  the  word  itself.  If  developed  in  intensity,  it 
impels  toward  vigorous  movement.  Every  vigor- 
ous movement  of  the  body  .  .  .  involves  also 
the  less  noticeable  cooperation  of  the  viscera,  es- 
pecially of  the  circulatory  and  respiratory.  The 
extra  demand  made  upon  the  muscles  that  move 
the  frame  involves  a  heightened  action  of  the 
nutrient  organs  which  supply  to  the  muscles  the 
material  for  their  energy."  The  researches  here 
reported  have  revealed  a  number  of  unsuspected 
ways  in  which  muscular  action  is  made  more  effi- 
cient because  of  emotional  disturbances  of  the 
viscera.  Every  one  of  the  visceral  changes  that 
have  been  noted — the  cessation  of  processes  in  the 
alimentary  canal  (thus  freeing  the  energy  supply 
for  other  parts) ;  the  shifting  of  blood  from  the 

215 


216  BODILY   CHANGES 

abdominal  organs,  whose  activities  are  deferable, 
to  the  organs  immediately  essential  to  muscular 
exertion  (the  lungs,  the  heart,  the  central  nervous 
system) ;  the  increased  vigor  of  contraction  of  the 
heart ;  the  quick  abolition  of  the  effects  of  muscu- 
lar fatigue ;  the  mobilizing  of  energy-giving  sugar 
in  the  circulation — every  one  of  these  visceral 
changes  is  directly  serviceable  in  making  the  or- 
ganism more  effective  in  the  violent  display  of 
energy  which  fear  or  rage  or  pain  may  involve. 

"Reservoirs  op  Power" 

That  the  major  emotions  have  an  energizing 
effect  has  been  commonly  recognized.*  Darwin 
testified  to  having  heard,  "as  a  proof  of  the 
exciting  nature  of  anger,  that  a  man  when  ex- 
cessively jaded  will  sometimes  invent  imaginary 
offences  and  put  himself  into  a  passion,  uncon- 
sciously for  the  sake  of  reinvigorating  him- 
self ;  and,"  Darwin  ^  continues,  "since  hearing 
this  remark,  I  have  occasionally  recognized  its  full 
truth."  Under  the  impulse  of  fear  also,  men  have 
been  known  to  achieve  extraordinary  feats  of 
running  and  leaping.     McDougall  ^  cites  the  in- 

*  Eussell  (The  Pima  Indians,  United  States  Bureau  of 
Ethnology,  1908,  p.  243)  relates  a  tale  told  by  the  Indians  to 
their  children,  in  which  an  injured  coyote  was  chasing  some 
quails.  "Finally  the  quails  got  tired,"  according  to  the 
story,  "but  the  coyote  did  not,  for  he  was  angry  and  did  not 
feel  fatigue." 


ENERGIZING   INFLUENCE  217 

stance  of  an  athlete  who,  when  pursued  as  a  boy 
by  a  savage  animal,  leaped  over  a  wall  which  he 
could  not  again  "clear"  until  he  attained  his  full 
stature  and  strength.  The  very  unusual  abilities, 
both  physical  and  mental,  which  men  have  exhib- 
ited in  times  of  stress  were  dealt  with  from  the 
psychological  point  of  view  by  William  James  * 
in  one  of  his  last  essays.  He  suggested  that  in 
every  person  there  are  "reservoirs  of  power" 
which  are  not  ordinarily  called  upon,  but  which 
are  nevertheless  ready  to  pour  forth  streams  of 
energy  if  only  the  occasion  presents  itself.  These 
figurative  expressions  of  the  psychologist  receive 
definite  and  concrete  exemplification,  so  far  as  the 
physical  exhibitions  of  power  are  concerned,  in 
the  highly  serviceable  bodily  changes  which  have 
been  described  in  the  foregoing  chapters. 

It  would  doubtless  be  incorrect  to  attempt  to 
account  for  all  the  increased  strength  and  tireless 
endurance,  which  may  be  experienced  in  periods 
of  great  excitement,  on  the  basis  of  abundant  sup- 
plies provided  then  for  muscular  contraction,  and 
a  special  secretion  for  avoiding  or  abolishing  the 
depressive  influences  of  fatigue.  Tremors,  mus- 
cular twitchings,  the  assumption  of  characteristic 
attitudes,  all  indicate  that  there  is  an  immensely 
augmented  activity  of  the  nervous  system — an  ac- 
tivity that  discharges  powerfully  even  into  parts 
not  directly  concerned  in  struggle,  as,  for  exam- 


218  BODILY   CHANGES 

pie,  into  the  muscles  of  voice,  causing  peculiar 
cries  or  warning  notes;  into  the  muscles  of  the 
ears,  drawing  them  back  or  causing  them  to  stand 
erect,  and  into  the  small  muscles  about  the  lips, 
tightening  them  and  revealing  the  teeth.  The 
typical  appearances  of  human  beings,  as  well  as 
lower  animals,  when  in  the  grip  of  such  deeply- 
agitating  emotions  as  fear  and  rage,  are  so  well 
recognized  as  to  constitute  a  primitive  and  com- 
mon means  of  judging  the  nature  of  the  experience 
through  which  the  organism  is  passing.  This  "pat- 
tern" response  of  the  nervous  system  to  an  emo- 
tion-provoking object  or  situation  is  probably 
capable  of  bringing  into  action  a  much  greater 
number  of  neurones  in  the  central  nervous  system 
than  are  likely  to  be  concerned  in  even  a  supreme 
act  of  volition.  The  nervous  impulses  delivered  to 
the  muscles,  furthermore,  operate  upon  organs 
well  supplied  with  energy-yielding  material  and 
well  fortified  by  rapidly  circulating  blood  and  by 
secreted  adrenin,  against  quick  loss  of  power  be- 
cause of  accumulating  waste.  Under  such  circum- 
stances of  excitement  the  performance  of  extraor- 
dinary feats  of  strength  or  endurance  is  natural 
enough.* 

*  If  individual  neurones  obey  the  law  of  either  supreme  ac- 
tion or  inaction,  the  "all-or-none  law,"  the  only  means  of 
securing  a  graded  response  is  through  variation  of  the  number 
of  neurones  engaged  in  action — the  more,  the  greater  the  re- 
sulting manifestation  of  strength. 


ENERGIZING   INFLUENCE  219 

In  connection  with  the  conception  that  strong 
emotion  has  a  dynamogenic  value,  it  is  of  interest 
to  note  that  on  occasions  when  great  demands  are 
likely  to  be  placed  on  the  neuro-muscular  system  in 
the  doing  of  unusual  labors,  emotional  excitement 
is  not  uncommonly  an  accompaniment.  In  order 
to  emphasize  points  in  the  argument  developed 
thus  far,  I  propose  to  cite  some  examples  of  the 
association  of  emotional  excitement  with  remark- 
able exhibitions  of  power  or  resistance  to  fatigue. 

The  Excitements  and  Energies  of  Competitive  Sports 

Already  in  an  earlier  account  (see  p.  75)  I  have 
mentioned  finding  sugar  in  the  urine  in  approxi- 
mately fifty  per  cent  of  a  group  of  college  football 
players  after  the  most  exacting  game  of  the  sea- 
son's play.  As  is  well  understood,  such  games  are 
heralded  far  and  wide,  loyal  supporters  of  each 
college  may  travel  hundreds  of  miles  to  attend  the 
contest,  enthusiastic  meetings  of  undergraduate 
students  are  held  in  each  college  to  demonstrate 
their  devotion  to  the  team  and  their  confidence  in 
its  prowess — indeed,  the  arguments  for  victory, 
the  songs,  the  cheering,  are  likely  to  be  so  disturb- 
ing to  the  players,  that  before  an  important  con- 
test they  are  not  infrequently  removed  from 
college  surroundings  in  order  to  avoid  being  over- 
wrought when  the  contest  comes. 

On  the  day  of  the  contest  the  excitement  is  mul- 


220  BODILY   CHANGES 

tiplied  manyfold.  There  is  practically  a  holiday 
in  college  and  to  a  large  extent  in  the  city  as  well. 
The  streets  are  filled  with  eager  supporters  of 
each  team  as  the  hosts  begin  to  gather  at  the  field. 
As  many  as  70,000  spectators  may  be  present,  each 
one  tense  and  strongly  partisan.  The  student 
bands  lead  the  singing,  by  thousands  of  voices, 
of  songs  which  urge  to  the  utmost  effort  for  the 
college ;  and,  in  anticipation,  these  songs  also  cele- 
brate the  victory. 

Into  the  midst  of  that  huge,  cheering,  yelling, 
singing,  flag-waving  crowd,  the  players  are  wel- 
comed in  a  special  outburst  of  these  same  demon- 
strations of  enthusiasm.  Soon  the  game  begins. 
The  position  of  every  player  is  known,  if  not  be- 
cause of  previous  acquaintance  and  recognition, 
because  card-diagrams  give  the  information. 
Every  important  play  is  seen  by  the  assembled 
thousands,  and  the  player  who  makes  it  is  at  once 
announced  to  all,  and  is  likely  to  be  honored  by  his 
multitudinous  college  mates  in  a  special  cheer, 
ending  in  his  name.  Any  player  who,  by  infrac- 
tion of  the  rules  or  failure  to  do  his  part,  loses 
ground  gained  by  his  team  is  also  known.  The  man 
who  is  "played  out"  in  efforts  to  win  for  his  team 
and  college,  and  consequently  has  to  leave  the  field, 
is  welcomed  to  the  side  lines  by  acclamations 
suited  for  a  great  hero.  In  short,  every  effort  is 
made,  through  the  powerful  incentives  of  censure 


ENERGIZING   INFLUENCE  221 

and  a  flaunting  recognition,  to  make  each  member 
of  the  team  realize  vividly  bis  responsibility,  both 
l^ersonal  and  as  one  of  a  group,  for  the  supreme, 
all-important  result — victory  for  his  college. 

This  responsibility  works  tremendously  on  the 
emotions  of  the  players.  In  the  dressing  room 
before  a  critical  contest  I  have  seen  a  "gridiron 
warrior,"  ready  in  canvas  suit,  cleated  shoes,  and 
leather  helmet,  sitting  grimly  on  a  bench,  his  fists 
clenched,  his  jaws  tight,  and  his  face  the  color  of 
clay.  He  performed  wonderfully  when  the  game 
began,  and  after  it  was  over  there  was  a  large  per- 
centage of  sugar  in  his  urine !  Probably  no  sport 
requires  a  more  sustained  and  extreme  display  of 
neuro-muscular  effort  than  American  football. 
And  from  the  foregoing  description  of  the  condi- 
tions that  surround  the  contests  it  is  easy  to  real- 
ize that  they  conspire  to  arouse  in  the  players  ex- 
citements which  would  bring  forth  very  efficiently 
the  bodily  reserves  for  use  in  the  fierce  struggle 
which  the  game  requires. 

What  is  true  of  football  is  true,  though  perhaps 
to  a  less  degree,  of  the  racing  sports,  as  running 
and  rowing.  Again  great  multitudes  attend  the 
events,  the  contests  are  followed  closely  from  be- 
ginning to  end,  and  as  the  goal  is  approached  the 
cheering  and  cries  for  victory  gather  in  volume 
and  intensity  as  if  arranged  for  a  thrilling  climax. 
The  whole  setting  is  most  highly  favorable  to  the 


222  BODILY   CHANGES 

dramatic  development  of  an  acme  of  excitement 
as  the  moment  for  the  last  desperate  effort  to  win 
is  pnt  forth. 

Frenzy  and  Endurance  in  Ceremonial  and  Other  Dances 

Dancing,  which  formed  a  significant  feature  of 
primitive  rituals,  has  always  been  accompanied  by- 
exciting  conditions,  and  not  unusually  was  an  ex- 
hibition of  remarkable  endurance.  In  the  trans- 
fer of  the  Ark  to  Zion  there  were  processions  and 
sacrifices,  and  King  David  "danced  before  the 
Lord  with  all  his  might."  Mooney  ^  in  his  ac- 
count of  dances  among  the  American  Indians  tells 
of  a  young  man  who  in  one  of  the  ceremonials 
danced  three  days  and  nights  without  food,  drink 
or  sleep.  In  such  a  terrible  ordeal  the  favoring 
presence  of  others,  who  through  group  action  help 
to  stimulate  both  the  excitement  and  the  activities, 
must  be  an  important  element  in  prolonging  the 
efforts  of  the  individual. 

In  the  history  of  religious  manias  ®  there  are 
many  instances  of  large  numbers  of  people  becom- 
ing frenzied  and  then  showing  extraordinary  en- 
durance while  dancing.  In*  1374  a  mania  broke 
forth  in  Germany,  the  Netherlands  and  France,  in 
which  the  victims  claimed  to  dance  in  honor  of 
Saint  John.  Men  and  women  went  about  dancing 
hand  in  hand,  in  pairs,  or  in  a  circle,  on  the  streets, 
in  the  churches,  at  their  homes,  or  wherever  they 


ENERGIZING   INFLUENCE  223 

miglit  be,  hour  after  hour  without  rest.  AVhile 
dancing  they  sang,  uttered  cries,  and  saw  visions. 
Whole  companies  of  .these  crazy  fanatics  went 
dancing  along  the  public  roads  and  into  the  cities, 
until  they  had  to  be  interfered  with. 

In  1740  an  extraordinary  sect,  known  as  the 
"Jumpers,"  arose  in  Wales.  According  to  the 
description  given  by  Wesley,  their  exercises  were 
not  unlike  those  of  certain  frenzied  states  among 
the  Indians.  "After  the  preaching  was  over," 
Wesley  '^  wrote,  "anyone  who  pleased  gave  out 
a  verse  of  a  hjTnn;  and  this  they  sung  over  and 
over  again,  with  all  their  might  and  main,  thirty 
or  forty  times,  till  some  of  them  worked  them- 
selves into  a  sort  of  drunkenness  or  madness ;  they 
were  then  violently  agitated,  and  leaped  up  and 
down  in  all  manner  of  postures,  frequently  for 
hours  together."  There  were  sometimes  thou- 
sands at  a  single  meeting  of  the  Jumpers,  shouting 
out  their  excitement  and  ready  to  leap  for  joy.^ 
Wesley  has  also  described  instances  of  tremendous 
emotional  outburst  at  Methodist  meetings  which 
he  addressed.  "Some  were  torn  with  a  kind  of 
convulsive  motion  in  every  part  of  their  bodies, 
and  that  so  violently  that  often  four  or  five  per- 
sons could  not  hold  one  of  them.  I  have  seen 
many  hysterical  or  epileptic  fits,"  he  wrote,  "but 
none  of  them  were  like  these  in  many  respects." 

Among  the  dervishes  ^  likewise  the  dance  is  ac- 


224  BODILY   CHANGES 

companied  by  intense  excitement  and  apparently 
tireless  movements.  "The  cries  of  'Ya  Allah !'  are 
increased  doubly,  as  also  those  of  'Ya  Hoo !'  with 
frightful  bowlings  shrieked  by  the  dervishes  to- 
gether in  the  dance."  .  .  .  "There  was  no  reg- 
ularity in  their  dancing,  but  each  seemed  to  be  per- 
forming the  antics  of  a  madman ;  now  moving  his 
body  up  and  down;  the  next  moment  turning 
round,  then  using  odd  gesticulations  with  his  arms, 
next  jumping,  and  sometimes  screaming."  .  .  . 
"At  the  moment  when  they  would  seem  to  stop 
from  sheer  exhaustion  the  sheikh  makes  a  point 
of  exciting  them  to  new  efforts  by  walking  through 
their  midst,  making  also  himself  most  violent 
movements.  He  is  next  replaced  by  two  elders, 
who  double  the  quickness  of  the  step  and  the  agita- 
tion of  the  body ;  they  even  straighten  themselves 
up  from  time  to  time,  and  excite  the  envy  or  emu- 
lation of  others  in  their  astonishing  efforts  to  con- 
tinue the  dance  until  their  strength  is  entirely 
exhausted."  Such  is  the  frenzy  thus  developed 
that  the  performers  may  be  subjected  to  severe 
pain,  yet  only  show  signs  of  elation. 

In  all  these  dances  the  two  most  marked  features 
are  the  intense  excitement  of  those  who  engage  in 
them  and  the  very  remarkable  physical  endurance 
which  they  manifest.  Although  there  is  no  direct 
evidence,  such  as  was  obtained  in  examining  the 
football  players,  that  bodily  changes  favorable  to 


ENERGIZIXa   INFLUENCE  225 

great  neuro-miiscular  exertion  are  developed  in 
these  fnries  of  fanaticism,  it  is  highly  probable 
that  they  are  so  developed,  and  that  the  feats  of 
fortitude  which  are  performed  are  to  a  large  ex- 
tent explicable  on  the  basis  of  a  "tapping  of  the 
reservoirs  of  power"  through  the  emotional  ex- 
citement. 

The  Fierce  Emotions  and  Struggles  of  Battle 

Throughout  the  discussion  of  the  probable  sig- 
nificance of  the  bodily  changes  in  pain  and  great 
emotion,  the  value  of  these  changes  in  the  strug- 
gles of  conflict  or  escape  was  emphasized.  In  hu- 
man beings  as  well  as  in  lower  animals  the  wildest 
passions  are  aroused  when  the  necessities  of  com- 
bat become  urgent.  One  needs  only  to  glance  at 
the  history  of  warfare  to  observe  that  when  the 
primitive  emotions  of  anger  and  hatred  are  per- 
mitted full  sway,  men  who  have  been  considerate 
and  thoughtful  of  their  fellows  and  their  fellows' 
rights  suddenly  may  turn  into  infuriated  savages, 
slaughtering  innocent  women  and  children,  muti- 
lating the  wounded,  burning,  ravaging,  and  looting, 
with  all  the  wild  fervor  of  demons.  It  is  in  such 
excesses  of  emotional  turbulence  that  the  most 
astonishing  instances  of  prolonged  exertion  and 
incredible  endurance  are  to  be  found. 

Probably  the  fiercest  struggles  between  men 
that  are  recorded  are  those  which  occurred  when 


226  BODILY   CHANGES 

the  wager  of  battle  was  a  means  of  determining 
innocence  or  guilt.    In  the  corners  of  the  plot  se- 
lected for  the  combat  a  bier  was  prepared  for  each 
participant,  as  a  symbol  that  the  struggle  was  for 
life  or  death.    Each  was  attended  by  his  relatives 
and   followers,    and   by   his    father    confessor.^^ 
After  each  had  prayed  to  God  for  help  in  the  com- 
ing combat,  the  weapons  were  selected,  the  sacra- 
ment was  administered,  and  the  battle  was  begun. 
The  principals  fought  to  the  end  with  continuous 
and  brutal  ferocity,  resembling  the  desperate  en- 
counters of  wild  beasts.    A  fairly  illustrative  ex- 
ample is  furnished  in  an  incident  which  followed 
the  assassination  of  Charles  the  Good  of  Flanders 
in  1127.    One  of  the  accomplices,  a  knight  named 
Guy,  was  challenged  for  complicity  by  another 
named  Herman.     Both  were  renowned  warriors. 
Herman  was  speedily  unhorsed  by  Guy,  who  with 
his  lance  frustrated  all  Herman's  attempts  to  re- 
mount.   Then  Herman  disabled  Guy's  horse,  and 
the  combat  was  renewed  on  foot  with  swords. 
Equally  skilful  in  fence,  they  continued  the  strug- 
gle till  fatigue  compelled  them  to  drop  sword  and 
shield,  whereupon  they  wrestled  for  the  mastery. 
Guy  threw  his  antagonist,  fell  on  him,  and  beat 
him  in  the  face  with  his  gauntlets  till  he  seemed  to 
be  motionless ;  but  Herman  had  quietly  slipped  his 
hand  below  the  other's  coat  of  mail  and,  grasping 
the  testicles,  with  a  mighty  effort  wrenched  them 


ENERGIZING   INFLUENCE  227 

away.  Immediately  Guy  fell  over  and  expired. ^^ 
In  such  terrific  fights  as  these,  conducted  in  the 
extremes  of  rage  and  hate,  the  mechanisms  for 
reenforcing  the  parts  of  the  body  which  are  of 
primary  importance  in  the  struggle  are  brought 
fully  into  action  and  are  of  utmost  value  in  secur- 
ing victory. 

The  Stimulating  Influence  of  Witnesses  and  of  Music 

It  is  noteworthy  that  in  all  the  instances  thus 
far  cited — in  the  great  games,  in  dancing,  and  in 
fighting — two  factors  are  present  that  are  well 
known  to  have  an  augmenting  effect  both  in  the 
full  development  of  emotions  and  in  the  perform- 
ance of  unusual  muscular  labors.  One  of  these  is 
the  crowd  of  witnesses  or  participants,  who  con- 
tribute the  "mob  spirit"  that  tends  to  carry  the 
actions  of  the  individual  far  beyond  the  limits  set 
by  any  personal  considerations  or  prudencies. 
The  other  is  the  influence  of  music.  As  Darwin 
long  ago  indicated,  music  has  a  wonderful  power 
of  recalling  in  a  vague  and  indefinite  manner 
strong  emotions  which  have  been  felt  by  our  an- 
cestors in  long-past  ages.  Especially  is  this  true 
of  martial  music.  For  the  grim  purposes  of  war 
the  reed  and  the  lute  are  grotesquely  ill-suited;  to 
rouse  men  to  action  strident  brass  and  the  jarring 
instruments  of  percussion  are  used  in  full  force. 
The  influence  of  martial  music  on  some  persons 


228  BODILY   CHANGES 

is  so  profound  as  to  cause  the  muscles  to  tremble 
and  tears  to  come  to  the  eyes— both  indications  of 
the  deep  stirring  of  emotional  responses  in  the 
body.  And  when  deeds  of  fortitude  and  fierce  ex- 
ertion are  to  be  performed  the  effectiveness-  of 
such  music  in  rousing  the  aggressive  emotions  has 
long  been  recognized.  The  Eomans  charged  their 
foes  amid  the  blasts  of  trumpets  and  horns.  The 
ancient  Germans  rushed  to  battle,  their  forces 
spurred  by  the  sounds  of  drums,  flutes,  cymbals 
and  clarions.  There  is  a  tradition  that  the  Hunga- 
rian troops  are  the  worst  in  Europe,  until  their 
bands  begin  to  play — then  they  are  the  best !  The 
late  General  Linevitch  is  quoted  as  saying :  "Music 
is  one  of  the  most  vital  ammunitions  of  the  Russian 
army.  Without  music  a  Russian  soldier  would  be 
dull,  cowardly,  brutal  and  inefficient.  From  music 
he  absorbs  a  magic  power  of  endurance,  and  for- 
gets the  sufferings  and  mortality.  It  is  a  divine 
dynamite."  And  Napoleon  is  said  to  have  testified 
that  the  weird  and  barbaric  tunes  of  the  Cossack 
regiments  infuriated  them  to  such  rage  that  they 
wiped  out  the  cream  of  his  army.^^  A  careful 
consideration  of  the  use  of  martial  music  in  war- 
fare would  perhaps  bring  further  interesting  evi- 
dence that  its  function  is  to  reenforce  the  bodily 
changes  that  attend  the  belligerent  emotions. 

Only  a  few  instances  of  the  combination  of  ex- 
treme pain,  rage,  terror  or  excitement,  and  tre- 


ENEKGIZING    INFLUENCE  229 

mendous  muscular  power  have  been  given  in  the 
preceding  pages.  Doubtless  in  numerous  other 
conditions  these  two  groups  of  phenomena  occur 
together.  In  the  lives  of  firemen  and  the  police,  in 
the  experiences  of  escaping  prisoners,  of  ship- 
wrecked sailors,  in  the  struggles  between  pioneers 
and  their  savage  enemies,  in  accounts  of  forced 
marches  .or  retreats,  search  would  reveal  many  ex- 
amples of  such  bodily  disturbances  as  have  been 
described  in  earlier  chapters  as  augmenting  the 
effectiveness  of  muscular  efforts,  and  such  exhibi- 
tions of  power  or  endurance  as  are  evidently  far 
beyond  the  ordinary.  There  is  every  reason  for 
believing  that,  were  the  conditions  favorable  to  ex- 
perimental testing,  it  would  be  possible  to  demon- 
strate and  perhaps  to  measure  the  addition  to  the 
dynamics  of  bodily  action  that  appears  as  the  ac- 
companiment of  violent  emotional  disturbance. 

The  Feeling  of  Power 

In  this  connection  it  is  highly  significant  that  in 
times  of  strong  excitement  there  is  not  infrequent 
testimony  to  a  sense  of  overwhelming  power  that 
sweeps  in  like  a  sudden  tide  and  lifts  the  person 
to  a  new  high  level  of  ability.  A  friend  of  mine, 
whose  nature  is  somewhat  choleric,  has  told  me 
that  when  he  is  seized  with  anger,  he  is  also  pos- 
sessed by  an  intense  conviction  that  he  could  crush 
and  utterly  destroy  the  object  of  his  hostility.   And 


230  BODILY    CHANGES 

I  have  heard  a  football  player  confess  that  just 
before  the  final  game  such  an  access  of  strength 
seemed  to  come  to  him  that  he  felt  able,  on  the 
signal,  to  crouch  and  with  a  jump  go  crashing 
through  any  ordinary  door.  There  is  intense  sat- 
isfaction in  these  moments  of  supreme  elation, 
when  the  body  is  at  its  acme  of  accomplishment. 
And  it  is  altogether  probable  that  the  critical  dan- 
gers of  adventure  have  a  fascination  because  fear 
is  thrilling,  and  extrication  from  a  predicament,  by 
calling  forth  all  the  bodily  resources  and  setting 
them  to  meet  the  challenge  of  the  difficulty,  yields 
many  of  the  joys  of  conquest.  For  these  reasons 
vigorous  men  go  forth  to  seek  dangers  and  to  run 
large  chances  of  serious  injury.  "Danger  makes 
us  more  alive.  We  so  love  to  strive  that  we  come 
to  love  the  fear  that  gives  us  strength  for  conflict. 
Fear  is  not  only  something  to  be  escaped  from  to 
a  place  or  state  of  safety,  but  welcomed  as  an  ar- 
senal of  augmented  strength." ^^  And  thus  in 
the  hazardous  sports,  in  mountain  climbing,  in  the 
hunting  of  big  game,  and  in  the  tremendous  ad- 
venture of  war,  risks  and  excitement  and  the  sense 
of  power  surge  up  together,  setting  free  unsus- 
pected energies,  and  bringing  vividly  to  conscious- 
ness memorable  fresh  revelations  of  the  possibili- 
ties of  achievement. 


ENERGIZING   INFLUENCE  231 

EEFEKENCES 

^  Sherrington :  The  Integrative  Action  of  the  Nervous 
System,  New  York,  1906,  p.  265. 

-  Darwin :  The  Expression  of  Emotions  in  Man  and  Ani- 
mals, New  York,  1905,  p.  Y9. 

^  McDougall :  Introduction  to  Social  Psychology,  London, 
1908,  p.  50. 

*  James :  The  Energies  of  Men,  p.  227,  in  Memories  and 
Studies,  New  York,  1911. 

^  Mooney :  The  Ghost-Dance  Religion,  United  States 
Bureau  of  Ethnology,  1892-3,  p.  924. 

^SchafP:  Religious  Encyclopedia,  New  York,  1908,  iii, 
p.  346. 

^Southey:  Life  of  Charles  Wesley,  New  York,  1820,  ii, 
p.  164. 

^  Southey :  Loc.  cit.,  i,  p.  240. 

9  Brown :  The  Dervishes,  London,  1868,  pp.  218-222,  260. 

^°  Major:  Geschichte  der  Ordalien,  Jena,  1796,  pp.  258- 
261. 

^^  Lea :  Superstition  and  Force,  Philadelphia,  1892,  p.  178. 

'^Narodny:  Musical  America,  1914,  xx,  No.  14. 

^^  Hall :  American  Journal  of  Psychology,  1914,  xxv,  p. 
154. 


CHAPTER   XIII 

THE  NATURE  OF  HUNGER 

On  tlie  same  plane  with  pain  and  the  dominant 
emotions  of  fear  and  anger,  as  agencies  which  de- 
termine the  action  of  organisms,  is  the  sensation 
of  hunger.  It  is  a  sensation  so  peremptory,  so  dis- 
agreeable, so  tormenting,  that  men  have  commit- 
ted crimes  in  order  to  assuage  it.  It  has  led  to 
cannibalism,  even  among  the  civilized.  It  has  re- 
sulted in  suicide.  And  it  has  defeated  armies — 
for  the  aggressive  spirit  becomes  detached  from 
larger  loyalties  and  turns  personal  and  selfish  as 
hunger  pangs  increase  in  vigor  and  insistence. 

In  1905,  while  observing  in  myself  the  rhythmic 
sounds  produced  by  the  activities  of  the  alimentary 
tract,  I  had  occasion  to  note  that  the  sensation 
of  hunger  was  not  constant  but  recurrent,  and  that 
the  moment  of  its  disappearance  was  often  associ- 
ated with  a  rather  loud  gurgling  sound  as  heard 
through  the  stethoscope.  This  and  other  evidence, 
indicative  of  a  source  of  the  hunger  sensations  in 

232 


THE   NATURE    OF   HUNGER         233 

the  contractions  of  the  digestive  canal,  I  reported 
in  1911.^  That  same  year,  with  the  help  of  one 
of  my  students,  A.  L.  Washburn,  I  obtained  final 
proof  for  this  inference. 

Appetite  and  Hunger 

The  sensations  of  appetite  and  hunger  are  so 
complex  and  so  intimately  interrelated  that  any 
discussion  of  either  sensation  is  sure  to  go  astray 
unless  at  the  start  there  is  clear  understanding  of 
the  meanings  of  the  terms.  The  view  has  been 
propounded  that  appetite  is  the  first  degree  of 
hunger,  the  mild  and  pleasant  stage,  agreeable  in 
character;  and  that  hunger  itself  is  a  more  ad- 
vanced condition,  disagreeable  and  even  painful — 
the  unpleasant  result  of  not  satisfying  the  appe- 
tite.- On  this  basis  appetite  and  hunger  would 
differ  only  quantitatively.  Another  view,  which 
seems  more  justifiable,  is  that  the  two  experiences 
are  fundamentally  different. 

Careful  observation  indicates  that  appetite  is  re- 
lated to  previous  sensations  of  taste  and  smell  of 
food.  Delightful  or  disgusting  tastes  and  odors, 
associated  with  this  or  that  edible  substance,  de- 
termine the  appetite.  It  has,  therefore,  important 
psychic  elements  in  its  composition.  Thus,  by  tak- 
ing thought,  we  can  anticipate  the  odor  of  a  de- 
licious beefsteak  or  the  taste  of  peaches  and  cream, 
and  in  that  imagination  we  can  find  pleasure.    In 


234  BODILY   CHANGES 

the  realization,  direct  effects  in  the  senses  of  taste 
and  smell  give  still  further  delight.  As  already 
noted  in  the  first  chapter,  observations  on  experi- 
mental animals  and  on  human  beings  have  shown 
that  the  pleasures  of  both  anticipation  and  realiza- 
tion, by  stimulating  the  flow  of  saliva  and  gastric 
juice,  play  a  highly  significant  role  in  the  initiation 
of  digestive  processes. 

Among  prosperous  people,  supplied  with  abun- 
dance of  food,  the  appetite  seems  sufficient  to  en- 
sure for  bodily  needs  a  proper  supply  of  nutri- 
ment. We  eat  because  dinner  is  announced,  be- 
cause by  eating  we  avoid  unpleasant  consequences, 
and  because  food  is  placed  before  us  in  delectable 
form  and  with  tempting  tastes  and  odors.  Under 
less  easy  circumstances,  however,  the  body  needs 
are  supplied  through  the  much  stronger  and  more 
insistent  demands  of  hungery 

The  sensation  of  hunger  is  difficult  to  describe, 
but  almost  everyone  from  childhood  has  felt  at 
times  that  dull  ache  or  gnawing  pain  referred  to 
the  lower  mid-chest  region  and  the  epigastrium, 
which  may  take  imperious  control  of  human  ac- 
tions. As  Sternberg  has  pointed  out,  hunger  may 
be  sufficiently  insistent  to  force  the  taking  of  food 
which  is  so  distasteful  that  it  not  only  fails  to 
rouse  appetite,  but  may  even  produce  nausea.  The 
hungry  being  gulps  his  food  with  a  rush.  The 
pleasures  of  appetite  are  not  for  him — he  wants 


THE   NATURE    OF   HUNGER         235 

quantity  rather  than  quality,  and  he  wants  it  at 
once. 

Hunger  and  appetite  are,  therefore,  widely  dif- 
ferent— in  physiological  basis,  in  localization  and 
in  psychic  elements.  Hunger  may  be  satisfied 
while  the  appetite  still  calls.  Who  is  still  hungry 
when  the  tempting  dessert  is  served,  and  yet  are 
there  any  who  refuse  it,  on  the  plea  that  they  no 
longer  need  it!  On  the  other  hand,  appetite  may 
be  in  abeyance  while  hunger  is  goading.^  What 
ravenous  boy  is  critical  of  his  food  I  Do  we  not 
all  know  that  "hunger  is  the  best  sauce"  ?  Although 
the  two  sensations  may  thus  exist  separately,  they 
nevertheless  have  the  same  function  of  leading  to 
the  intake  of  food,  and  they  usually  appear  to- 
gether. Indeed,  the  cooperation  of  hunger  and  ap- 
petite is  probably  the  reason  for  their  being  so 
frequently  confused. 

The   Sensation  of  Hunger 

Hunger  may  be  described  as  having  a  central 
core  and  certain  more  or  less  variable  accessories. 
The  peculiar  dull  ache  of  hungriness,  referred  to 
the  epigastrium,  is  usually  the  organism's  first 
strong  demand  for  food ;  and  when  the  initial  or- 
der is  not  obeyed,  the  sensation  is  likely  to  grow 
into  a  highly  uncomfortable  pang  or  gnawing,  less 
definitely  localized  as  it  becomes  more  intense. 
This  may  be  regarded  as  the  essential  feature  of 


236  BODILY    CHANGES 

hunger.  Besides  the  dull  ache,  however,  lassitude 
and  drowsiness  may  appear,  or  faintness,  or  vio- 
lent headache,  or  irritability  and  restlessness  such 
that  continuous  effort  in  ordinary  affairs  becomes 
increasingly  difficult.  That  these  states  differ 
much  with  individuals — ^headache  in  one  and  faint- 
ness in  another,  for  example — indicates  that  they 
do  not  constitute  the  central  fact  of  hunger,  but 
are  more  or  less  inconstant  accompaniments.  The 
"feeling  of  emptiness,"  which  has  been  mentioned 
as  an  important  element  of  the  experience,^  is 
an  inference  rather  than  a  distinct  datum  of  con- 
sciousness, and  can  likewise  be  eliminated  from 
further  consideration.  The  dull  pressing  sensa- 
tion is  left,  therefore,  as  the  constant  character- 
istic, the  central  fact,  to  be  examined  in  detail. 

Hunger  can  evidently  be  regarded  from  the 
psychological  point  of  view,  and  discussed  solely 
on  the  basis  of  introspection ;  or  it  can  be  studied 
with  reference  to  its  antecedents  and  to  the  physi- 
ological conditions  which  accompany  it — a  consid- 
eration which  requires  the  use  of  both  objective 
methods  and  subjective  observation.  This  psycho- 
physiological treatment  of  the  subject  will  be  de- 
ferred till  the  last.  Certain  theories  which  have 
been  advanced  with  regard  to  hunger,  and  which 
have  been  given  more  or  less  credit,  must  first  be 
examined. 

Two  main  theories  have  been  advocated.    The 


THE   NATURE    OF   HUNGER         237 

first  is  supported  by  contentions  that  hunger  is  a 
general  sensation,  arising  at  no  special  region  of 
the  body,  but  having  a  local  reference.  This  the- 
ory has  been  more  widely  credited  by  physiologists 
and  psychologists  than  the  other.  The  other  is 
supported  by  evidence  that  hunger  has  a  local 
source  and  therefore  a  local  reference.  In  the 
course  of  our  examination  of  these  views  we  shall 
have  opportunity  to  consider  some  pertinent  new 
observations. 

The  Theory  that  Hunger  is  a  General  Sensation 

The  conception  that  hunger  arises  from  a  gen- 
eral condition  of  the  body  rests  in  turn  on  the  no- 
tion that,  as  the  body  uses  up  material,  the  blood 
becomes  impoverished.  Schiff  ^  advocated  this 
notion,  and  suggested  that  poverty  of  the  blood  in 
food  substance  affects  the  tissues  in  such  manner 
that  they  demand  a  new  supply.  The  nerve  cells 
of  the  brain  share  in  this  general  shortage  of  pro- 
visions, and  because  of  internal  changes,  give  rise 
to  the  sensation.  Thus  is  hunger  explained  as  an 
experience  dependent  on  the  body  as  a  whole. 

Three  classes  of  evidence  are  cited  in  support 
of  this  view: 

1.  "Hunger  increases  as  time  passes" — a  partial 
statement.  The  development  of  hunger  as  time 
passes  is  a  common  observation  which  quite  ac- 
cords with  the  assumption  that  the  condition  of  the 


238  BODILY   CHANGES 

body  and  the  state  of  the  blood  are  becoming  con- 
stantly worse,  so  long  as  the  need,  once  estab- 
lished, is  not  satisfied. 

While  it  is  true  that  with  the  lapse  of  time  hun- 
ger increases  as  the  supply  of  body  nutriment  de- 
creases, this  concomitance  is  not  proof  that  the 
sensation  arises  directly  from  a  serious  encroach- 
ment on  the  store  of  food  materials.  If  this  argu- 
ment were  valid  we  should  expect  hunger  to  become 
more  and  more  distressing  until  death  follows 
from  starvation.  There  is  abundant  evidence  that 
the  sensation  is  not  thus  intensified;  on  the  con- 
trary, during  continued  fasting  hunger,  at  least  in 
some  persons,  wholly  disappears  after  the  first  few 
days.  Luciani,^  who  carefully  recorded  the  ex- 
perience of  the  faster  Succi,  states  that  after  a  cer- 
tain time  the  hunger  feelings  vanish  and  do  not 
return.  And  he  tells  of  two  dogs  that  showed  no 
signs  of  hunger  after  the  third  or  fourth  day  of 
fasting;  thereafter  they  remained  quite  passive 
in  the  presence  of  food.  Tigerstedt,'^  who  also  has 
studied  the  metabolism  of  starvation,  declares  that 
although  the  desire  to  eat  is  very  great  during  the 
first  day  of  the  ordeal,  the  unpleasant  sensations 
disappear  early,  and  that  at  the  end  of  the  fast  the 
subject  may  have  to  force  himself  to  take  nourish- 
ment. The  subject,  "J,  A.,"  studied  by  Tigerstedt 
and  his  co-workers,^  reported  that  after  the  fourth 
day  of  fasting,  he  had  no  disagreeable  feelings. 


THE   NATUKE   OF   HUNGER         239 

Carrington,^  after  examining  many  persons 
who,  to  better  their  health,  abstained  from  eating 
for  different  periods,  records  that  "habit-hunger" 
usually  lasts  only  two  or  three  days  and,  if  plenty 
of  water  is  drunk,  does  not  last  longer  than  three 
days.  Viterbi,^"  a  Corsiean  lawyer  condemned 
to  death  for  political  causes,  determined  to  escape 
execution  by  depriving  his  body  of  food  and  drink. 
During  the  eighteen  days  that  he  lived  he  kept 
careful  notes.  On  the  third  day  the  sensation  of 
hunger  departed,  and  although  thereafter  thirst 
came  and  went,  hunger  never  returned.  Still  fur- 
ther evidence  of  the  same  character  could  be  cited, 
but  enough  has  already  been  given  to  show  that 
after  the  first  few  days  of  fasting  the  hunger  feel-, 
ings  may  wholly  cease.  On  the  theory  that  hunger 
is  a  manifestation  of  bodily  need,  are  we  to  sup- 
pose that,  in  the  course  of  starvation,  the  body  is 
mysteriously  not  in  need  after  the  third  day,  and 
that  therefore  the  sensation  of  hunger  disappears? 
The  absurdity  of  such  a  view  is  obvious. 

2.  "Hunger  may  be  felt  though  the  stomach  be 
full" — a  selected  alternative.  Instances  of  duo- 
denal fistula  in  man  have  been  carefully  studied, 
which  have  shown  that  a  modified  sensation  of 
hunger  may  be  felt  when  the  stomach  is  full.  A 
famous  case  described  by  Busch  ^^  has  been  re- 
peatedly used  as  evidence.  His  patient,  who  lost 
nutriment  through  a  duodenal  fistula,  was  hungry 


240  BODILY   CHANGES 

soon  after  eating,  and  felt  satisfied  only  when  the 
chyme  was  restored  to  the  intestine  through  the 
distal  fistulous  opening.  As  food  is  absorbed 
mainly  through  the  intestinal  wall,  the  inference 
is  direct  that  the  general  bodily  state,  and  not  the 
local  conditions  of  the  alimentary  canal,  must  ac- 
count for  the  patient's  feelings. 

A  full  consideration  of  the  evidence  from  cases 
of  duodenal  fistula  cannot  so  effectively  be  pre- 
sented now  as  later.  That  in  Busch's  case  hunger 
disappeared  while  food  was  being  taken  is,  as  we 
shall  see,  quite  significant.  It  may  be  that  the 
restoration  of  chyme  to  the  intestine  quieted 
hunger,  not  because  nutriment  was  thus  intro- 
duced into  the  body,  but  because  the  presence  of 
material  altered  the  nature  of  gastro-intestinal 
activity.  The  basis  for  this  suggestion  will  be 
given  in  due  course. 

3.  "Animals  may  eat  eagerly  after  section  of 
their  vagus  and  splanchnic  nerves" — a  fallacious 
argument.  The  third  support  for  the  view  that 
hunger  has  a  general  origin  in  the  body  is  derived 
from  observations  on  experimental  animals.  By 
severance  of  the  vagus  and  splanchnic  nerves,  the" 
lower  esophagus,  the  stomach  and  the  small  in- 
testine can  be  wholly  separated  from  the  central 
nervous  system.  Animals  thus  operated  upon 
nevertheless  eat  food  placed  before  them,  and  may 
indeed  manifest  some  eagerness  for  it.^^     How 


THE   NATURE   OF   HUNGER         241 

is  this  behavior  to  be  aecoimted  for — when  the 
possibility  of  local  stimulation  has  been  eliminated 
— save  by  assuming  a  central  origin  of  the  impulse 
to  eat? 

The  fallacy  of  this  evidence,  though  repeatedly 
overlooked,  is  easily  shown.  We  have  already  seen 
that  appetite  as  well  as  hunger  may  lead  to  the 
taking  of  food.  Indeed,  the  animal  with  all  gas- 
tro-intestinal  nerves  cut  may  have  the  same  in- 
centive to  eat  that  a  well-fed  man  may  have,  who 
delights  in  the  pleasurable  taste  and  smell  of  food 
and  knows  nothing  of  hunger  pangs.  Even  when 
the  nerves  of  taste  are  cut,  as  they  were  in 
Longet's  experiments,^^  sensations  of  smell  are 
still  possible,  as  well  as  agreeable  associations 
which  can  be  roused  by  sight.  More  than  fifty 
years  ago  Ludwig  ^^  pointed  out  that,  even  if 
all  the  nerves  were  severed,  psychic  reasons  could 
be  given  for  the  taking  of  food,  and  yet  because 
animals  eat  after  one  or  another  set  of  nerves  is 
eliminated,  the  conclusion  has  been  drawn  by  vari- 
ous writers  that  the  nerves  in  question  are  thereby 
proved  to  be  not  concerned  in  the  sensation  of 
hunger.  Evidently,  since  hunger  is  not  required 
for  eating,  the  act  of  eating  is  no  testimony  what- 
ever that  the  animal  is  hungry,  and,  after  the 
nerves  have  been  severed,  is  no  proof  that  hunger 
is  of  central  origin. 


242  BODILY   CHANGES 

.^^EAKNESS   OF    THE    ASSUMPTIONS   UNDERLYING   THE    ThEORY 

THAT  Hunger  is  a  General  Sensation 

The  evidence  thus  far  examined  has  been  shown 
to  afford  only  shaky  support  for  the  theory  that 
hunger  is  a  general  sensation.  The  theory,  fur- 
thermore, is  weak  in  its  fundamental  assumptions. 
There  is  no  clear  indication,  for  example,  that  the 
blood  undergoes  or  has  undergone  any  marked 
change,  chemical  or  physical,  when  the  first  stages 
of  hunger  appear.  There  is  no  evidence  of  any 
direct  chemical  stimulation  of  the  gray  matter  of 
the  cerebral  cortex.  Indeed,  attempts  to  excite  the 
gray  matter  artificially  by  chemical  agents  have 
been  without  results  ;^^  and  even  electrical  stim- 
ulation, which  is  effective,  must,  in  order  to 
produce  movements,  be  so  powerful  that  the  move- 
ments have  been  attributed  to  excitation  of  under- 
lying white  matter  rather  than  cells  in  the  gray. 
This  insensitivity  of  cortical  cells  to  direct  stimu- 
lation is  not  at  all  favorable  to  the  notion  that  they 
are  sentinels  set  to  warn  against  too  great  diminu- 
tion of  bodily  supplies. 

Body  Need  May  Exist  Without  Hunger 

Still  further  evidence  opposed  to  the  theory  that 
hunger  results  directly  from  the  using  up  of  or- 
ganic stores  is  found  in  patients  suffering  from 
fever.  Metabolism  in  fever  patients  is  augmented, 
body  substance  is  destroyed  to  such  a  degree  that 


THE   NATURE   OF   HUNGER         243 

the  weight  of  the  patient  may  be  greatly  reduced, 
and  yet  the  sensation  of  hunger  under  these  condi- 
tions of  increased  need  is  wholly  lacking. 

Again,  if  a  person  is  hungry  and  takes  food, 
the  sensation  is  suppressed  soon  afterwards,  long 
before  any  considerable  amount  of  nutriment 
could  be  digested  and  absorbed,  and  therefore 
long  before  the  blood  and  the  general  bodily  condi- 
tion, if  previously  altered,  could  be  restored  to 
normal. 

Furthermore,  persons  exposed  to  privation  have 
testified  that  hunger  can  be  temporarily  sup- 
pressed by  swallowing  indigestible  materials.  Cer- 
tainly scraps  of  leather  and  bits  of  moss,  not  to 
mention  clay  eaten  by  the  Otomacs,  would  not  ma- 
terially compensate  for  large  organic  losses.  In 
rebuttal  to  this  argument  the  comment  has  been 
made  that  central  states  as  a  rule  can  be  readily 
overwhelmed  by  peripheral  stimulation,  and  just 
as  sleep,  for  example,  can  be  abolished  by  bathing 
the  temples,  so  hunger  can  be  abolished  by  irritat- 
ing the  gastric  walls. ^'^  This  comment  is  beside 
the  point,  for  it  meets  the  issue  by  merely  assum- 
ing as  true  the  condition  under  discussion.  The 
absence  of  hunger  during  the  ravages  of  fever,  and 
its  quick  abolition  after  food  or  even  indigestible 
stuff  is  swallowed,  still  further  weakens  the  argu- 
ment, therefore,  that  the  sensation  arises  directly 
from  lack  of  nutriment  in  the  body. 


244  BODILY   CHANGES 

The  Theory  that  Hunger  is  of  General  Origin  Does  Not 

Explain  the  Quick  Onset  and  the  Periodicity 

of  the  Sensation 

Many  persons  have  noted  tliat  hunger  has  a 
sharp  onset.  A  person  may  be  tramping  in  the 
woods  or  working  in  the  fields,  where  fixed  atten- 
tion is  not  demanded,  and  without  premonition 
may  feel  the  abrupt  arrival  of  the  characteristic 
ache.  The  expression  "grub-struck"  is  a  pic- 
turesque description  of  this  experience.  If  this 
sudden  arrival  of  the  sensation  corresponds  to  the 
general  bodily  state,  the  change  in  the  general  bod- 
ily state  must  occur  with  like  suddenness  or  have 
a  critical  point  at  which  the  sensation  is  instantly 
precipitated.  There  is  no  evidence  whatever  that 
either  of  these  conditions  occurs  in  the  course  of 
metabolism. 

Another  peculiarity  of  hunger,  which  I  have  al- 
ready mentioned,  is  its  intermittency.  It  may 
come  and  go  several  times  in  the  course  of  a  few 
hours.  Furthermore,  while  the  sensation  is  pre- 
vailing, its  intensity  is  not  uniform,  but  marked 
by  ups  and  downs.  In  some  instances  the  ups  and 
downs  change  to  a  periodic  presence  and  absence 
without  change  of  rate.  In  my  own  experience  the 
hunger  pangs  came  and  went  on  one  occasion  as 
follows : 


Came 

Went 

—37—20 

38—30 

40—45 

41—10 

THE   NATURE   OF   HUNGER         245 

Came  Went 

41—45  42—25 

43—20  43—35 

44-^0  45—55 

46—15  46—30 

and  so  on,  for  ten  minutes  longer.  Again  in  this 
relation,  the  intermittent  and  periodic  character 
of  hunger  would  require,  on  the  theory  under  ex- 
amination, that  the  bodily  supplies  be  intermittent- 
ly and  periodically  insufficient.  During  one  mo- 
ment the  absence  of  hunger  would  imply  an 
abundance  of  nutriment  in  the  organism,  ten  sec- 
onds later  the  presence  of  hunger  would  imply 
that  the  stores  had  been  suddenly  reduced,  ten 
seconds  later  still  the  absence  of  hunger  would 
imply  a  sudden  renewal  of  plenty.  Such  zig-zag 
shifts  of  the  general  bodily  state  may  not  be  im- 
possible, but  from  all  that  is  known  of  the  course 
of  metabolism,  such  quick  changes  are  highly  im- 
probable. The  periodicity  of  hunger,  therefore,  is 
further  evidence  against  the  theory  that  the  sensa- 
tion has  a  general  basis  in  the  body. 

The  Theory  that  Hunger  is  of  General  Origen  Does  Not 
Explain  the  Local  Keference 

The  last  objection  to  this  theory  is  that  it  does 
not  account  for  the  most  common  feature  of 
hunger — namely,  the  reference  of  the  sensation  to 
the  region  of  the  stomach.  Schiff  and  others  ^'^ 
who  have   supported  the  theory  have  met  this 


246  BODILY   CHANGES 

objection  by  two  contentions.  First  tbey  have 
pointed  out  that  the  sensation  is  not  always  re- 
ferred to  the  stomach.  Schiff  interrogated  igno- 
rant soldiers  regarding  the  local  reference;  sev- 
eral indicated  the  neck  or  chest,  twenty-three  the 
sternum,  four  were  uncertain  of  any  region,  and 
two  only  designated  the  stomach.  In  other  words, 
the  stomach  region  was  most  rarely  mentioned. 

The  second  contention  against  the  importance 
of  local  reference  is  that  such  evidence  is  falla- 
cious. An  armless  man  may  feel  tinglings  which 
seem  to  arise  in  fingers  which  have  long  since 
ceased  to  be  a  portion  of  his  body.  The  fact  that 
he  experiences  such  tinglings  and  ascribes  them  to 
dissevered  parts,  does  not  prove  that  the  sensa- 
tion originates  in  those  parts.  And  similarly  the 
assignment  of  the  ache  of  hunger  to  any  special 
region  of  the  body  does  not  demonstrate  that  the 
ache  arises  from  that  region.  Such  are  the  argu- 
ments against  a  local  origin  of  hunger. 

Concerning  these  arguments  we  may  recall,  first, 
Schiff's  admission  that  the  soldiers  he  questioned 
were  too  few  to  give  conclusive  evidence.  Further, 
the  testimony  of  most  of  them  that  hunger  seemed 
to  originate  in  the  chest  or  region  of  the  sternum 
cannot  be  claimed  as  unfavorable  to  a  peripheral 
source  of  the  sensation.  The  description  of  feel- 
ings which  develop  from  disturbances  within  the 
body  is   almost  always   indefinite.     As   Head  ^^ 


THE   NATURE    OF   HUNGER         247 

and  others  have  shown,  conditions  in  a  viscus 
which  give  rise  to  sensation  are  likely  not  to  be  at- 
tributed to  the  viscus,  but  to  related  skin  areas. 
Under  such  circumstances  we  do  not  dismiss 
the  testimony  as  worthless  merely  because  it 
may  not  point  precisely  to  the  source  of  the 
trouble.  On  the  contrary,  we  use  such  testimony 
constantl}^  as  a  basis  for  judging  internal  dis- 
orders. 

With  regard  to  the  contention  that  reference  to 
the  periphery  is  not  proof  of  the  peripheral  origin 
of  a  sensation,  we  may  answer  that  the  force  of 
that  contention  depends  on  the  amount  of  acces- 
sory evidence  which  is  available.  Thus  if  we  see 
an  object  come  into  contact  with  a  finger,  we  are 
justified  in  assuming  that  the  simultaneous  sensa- 
tion of  touch  which  we  refer  to  that  finger  has  re- 
sulted from  the  contact,  and  is  not  a  purely  central 
experience  accidentally  attributed  to  an  outlying 
member.  Similarly  in  the  case  of  hunger — all  that 
we  need  as  support  for  the  peripheral  reference  of 
the  sensation  is  proof  that  conditions  occur  there, 
simultaneously  with  hunger  pangs,  which  might 
reasonably  be  regarded  as  giving  rise  to  those 
pangs. 

With  the  requirement  in  mind  that  peripheral 
conditions  be  adequate,  let  us  examine  the  state 
of  the  fasting  stomach  to  see  whether,  indeed,  con- 
ditions may  be  present  in  times  of  hunger  which 


248  BODILY   CHANGES 

would  sustain  the  theory  that  hunger  has  a  local 
outlying  source. 

Hunger  Not  Due  to  Emptiness  of  the  Stomach 

Among  the  suggestions  which  have  been  offered 
to  account  for  a  peripheral  origin  of  the  sensation 
is  that  of  attributing  it  to  emptiness  of  the  stom- 
ach. By  use  of 'the  stomach  tube  Nicolai  ^^  found 
that  when  his  subjects  had  their  first  intima- 
tion of  hunger  the  stomach  was  quite  empty. 
But,  in  other  instances,  after  lavage  of  the  stomach, 
the  sensation  did  not  appear  for  intervals  vary- 
ing between  one  and  a  half  and  three  and  a  half 
hours.  During  these  intervals  the  stomach  must 
have  been  empty,  and  yet  no  sensation  was  experi- 
enced. The  same  testimony  was  given  long  before 
by  Beaumont,-^  who,  from  his  observations  on 
Alexis  St.  Martin,  declared  that  hunger  arises  some 
time  after  the  stomach  is  normally  evacuated. 
Mere  emptiness  of  the  organ,  therefore,  does  not 
explain  the  phenomenon. 

Hunger  Not  Due  to  Hydrochloric  Acid  in  the  Empty 
Stomach 

A  second  theory,  apparently  suggested  by  obser- 
vations on  cases  of  hyperacidity,  is  that  the  ache  or 
pang  is  due  to  the  natural  hydrochloric  acid  of  the 
stomach  but  secreted  while  the  organ  is  empty. 
Again  the  facts  are  hostile.     Nicolai  ^^  reported 


THE   NATURE    OF   HUNGER         249 

that  the  gastric  wash-water  from  his  hungry  sub- 
jects was  neutral  or  only  slightly  acid.  This 
testimony  confirms  Beaumont's  statement,  and 
is  in  complete  agreement  with  the  results  of 
gastric  examination  of  fasting  animals  reported 
by  numerous  experimenters.  There  is  no  secre- 
tion into  the  empty  stomach  during  the  first  days 
of  starvation.  Furthermore,  persons  suffering 
from  absence  of  hydrochloric  acid  (acliylia  gas- 
trica)  declare  that  they  have  normal  feelings  of 
hunger.  Hydrochloric  acid  cannot,  therefore,  be 
called  upon  to  account  for  the  sensation. 

Hunger  Not  Due  to  Turgescence  of  the  Gastric  Mucous 
Membrane 

Another  theory,  which  was  first  advanced  by 
Beaumont,--  is  that  hunger  arises  from  turges- 
cence of  the  gastric  glands.  The  disappearance  of 
the  pangs  as  fasting  continues  has  been  accounted 
for  by  supposing  that  the  gastric  glands  share  in 
the  general  depletion  of  the  body,  and  that  thus  the 
turgescence  is  relieved.*  This  turgescence  theory 
has  commended  itself  to  several  recent  writers. 
Thus  Luciani  -^  has  accepted  it,  and  by  adding 
the  idea  that  nerves  distributed  to  the  mucosa  are 

*  A  better  explanation  perhaps  is  aflForded  by  Boldireff's 
discovery  that  at  the  end  of  two  or  three  days  the  stomachs 
of  fasting  dogs  begin  to  secrete  gastric  juice  and  continue 
the  secretion  indefinitely.  (BoldireflF,  Archives  Biologiques 
de  St.  Petersburg,  1905,  xi,  p.  98.) 


250  BODILY    CHANGES 

specially  sensitive  to  deprivation  of  food  he  ac- 
counts for  the  hunger  pangs.  Also  Valenti  ^^ 
declared  a  few  years  ago  that  the  turgescence 
theory  of  Beaumont  is  the  only  one  with  a  sem- 
blance of  truth  in  it.  The  experimental  work  re- 
ported by  these  two  investigators,  however,  does 
not  necessarily  sustain  the  turgescence  theory. 
Luciani  severed  the  previously  exposed  vagi  after 
cocainizing  them,  and  Valenti  merely  cocainized 
the  nerves;  the  fasting  dogs,  eager  to  eat  a  few 
minutes  previous  to  this  operation,  now  ran  about 
as  before,  but  when  offered  food,  licked  and  smelled 
it,  but  did  not  take  it.  This  total  neglect  of  the 
food  lasted  varying  periods  up  to  two  hours.  The 
vagus  nerves  seem,  indeed,  to  convey  impulses 
which  affect  the  procedure  of  eating,  but  there  is 
no  clear  evidence  that  those  impulses  arise  from 
distention  of  the  gland  cells.  The  turgescence 
theory,  moreover,  does  not  explain  the  effect  of 
taking  indigestible  material  into  the  stomach. 
According  to  Pawlow,  and  to  others  who  have  ob- 
served human  beings,  the  chewing  and  swallowing 
of  unappetizing  stuff  does  not  cause  any  secretion 
of  gastric  juice  (see  p.  8).  Yet  such  stuff  when 
swallowed  will  cause  the  disappearance  of  hunger, 
and  Nicolai  found  that  the  sensation  could  be  abol- 
ished by  simply  introducing  a  stomach  sound.  It 
is  highly  improbable  that  the  turgescence  of  the 
gastric  glands  can  be  reduced  by  either  of  these 


THE   NATURE   OF   HUNGER         251 

procedures.  The  turgescence  theory,  furthermore, 
does  not  explain  the  quick  onset  of  hunger,  or  its 
intermittent  and  periodic  character.  That  the  cells 
are  repeated!}^  swollen  and  contracted  within  peri- 
ods a  few  seconds  in  duration  is  almost  inconceiv- 
able. For  these  reasons,  therefore,  the  theory  that 
hunger  results  from  turgescence  of  the  gastric 
mucosa  can  reasonably  be  rejected. 

I  /  Hunger  the  Eesult   of  Contractions 

There  remain  to  be  considered,  as  a  possible 
cause  of  hunger-pangs,  contractions  of  the  stomach 
and  other  parts  of  the  alimentary  canal.  This  sug- 
gestion is  not  new.  Sixty-nine  years  ago  Weber  ^^ 
declared  his  belief  that  "strong  contraction  of 
the  muscle  fibres  of  the  wholly  empty  stomach, 
whereby  its  cavity  disappears,  makes  a  part  of  the 
sensation  which  we  call  hunger."  Vierordt  -"  drew 
the  same  inference  twenty-five  years  later  (in 
1871),  and  since  then  Ewald,  Knapp,  and  Hertz 
have  declared  their  adherence  to  this  view.  These 
writers  have  not  brought  forward  any  direct  evi- 
dence for  their  conclusion,  though  Hertz  has  cited 
Boldireff's  observations  on  fasting  dogs  as  prob- 
ably accounting  for  what  he  terms  "the  gastric 
constituent  of  the  sensation." 

The  Empty  Stomach  and  Intestine  Contract 

The  argument  commonly  used  against  the  gas- 
tric contraction  theory  is  that  the  stomach  is  not 


252  BODILY   CHANGES 

energetically  active  when  empty.  Thus  Schiff  ^^ 
stated,  "The  movements  of  the  empty  stomach 
are  rare  and  much  less  energetic  than  during  diges- 
tion." Luciani  ^^  expressed  his  disbelief  by  as- 
serting that  gastric  movements  are  much  more  ac- 
tive during  gastric  digestion  than  at  other  times, 
and  cease  almost  entirely  when  the  stomach  has 
discharged  its  contents.  And  Valenti^^  stated 
(1910),  "We  know  very  well  that  gastric  move- 
ments are  exaggerated  while  digestion  is  proceed- 
ing in  the  stomach,  but  when  the  organ  is  empty 
they  are  more  rare  and  much  less  pronounced," 
and,  therefore,  they  cannot  account  for  hunger. 

Evidence  opposed  to  these  suppositions  has  been 
in  existence  for  many  years.  In  1899  Bettmann  ^^ 
called  attention  to  the  contracted  condition  of 
the  stomach  after  several  days'  fast.  In  1902 
Wolff  ^^  reported  that  after  forty-eight  hours 
without  food  the  stomach  of  the  cat  may  be  so  small 
as  to  look  like  a  slightly  enlarged  duodenum.  In  a 
similar  circumstance  I  have  noticed  the  same  ex- 
traordinary smallness  of  the  organ,  especially  in 
the  pyloric  half.  The  anatomist  His  ^^  also  re- 
corded his  observation  of  the  phenomenon.  In  1905 
Boldireff^^  demonstrated  that  the  whole  gastro- 
intestinal tract  has  a  periodic  activity  while  not  di- 
gesting. Each  period  of  activity  lasts  from  twenty 
to  thirty  minutes,  and  is  characterized  in  the  stom- 
ach by  rhythmic  contractions  ten  to  twenty  in  num- 


THE   NATURE   OF   HUNGER         253 

ber.  These  contractions,  Boldireff  reports,  may  be 
stronger  than  during  digestion,  and  his  published 
records  clearly  support  this  statement.  The  inter- 
vals of  repose  between  periodic  recurrences  of  the 
contractions  lasted  from  one  and  a  half  to  two  and 
a  half  hours.  Especially  noteworthy  is  Boldireff's 
observation  that  if  fasting  is  continued  for  two  or 
three  days,  the  groups  of  contractions  appear  at 
gradually  longer  intervals  and  last  for  gradually 
shorter  periods,  and  thereupon,  as  the  gastric 
glands  begin  continuous  secretion,  all  movements 
cease. 

Observations  Suggesting  that  Contractions  Cause 
Hunger 

The  research,  previously  mentioned,  on  the 
rhythmic  sounds  produced  by  the  digestive  pro- 
cess, I  was  engaged  in  when  Boldireff's  paper  was 
published.  That  contractions  of  the  alimentary 
canal  on  a  gaseous  content  might  explain  the  hun- 
ger pangs  which  I  had  noticed  seemed  probable  at 
that  time,  especially  in  the  light  of  Boldireff's  ob- 
servations. Indeed,  Boldireff  ^^  himself  had  con- 
sidered hunger  in  relation  to  the  activities  he 
described,  but  solely  with  the  idea  that  hunger 
might  provoke  them;  and  since  the  activities  dwin- 
dled in  force  and  frequency  as  time  passed,  where- 
as, in  his  belief,  they  should  have  become  more  pro- 
nounced, he  a])andoned  the  notion  of  any  relation 


254  BODILY    CHANGES 

between  the  phenomena.  Did  not  Boldireff  misin- 
terpret his  own  observations?  When  he  was  con- 
sidering whether  hunger  might  cause  the  contrac- 
tions, did  he  not  overlook  the  possibility  that  the 
contractions  might  cause  hunger?  A  number  of 
experiences  have  led  to  the  conviction  that  Boldi- 
reff  did,  indeed,  fail  to  perceive  part  of  the  signifi- 
cance of  his  results.  For  example,  I  have  noticed 
the  disappearance  of  a  hunger  pang  as  gas  was 
heard  gurgling  upward  through  the  cardia.  That 
the  gas  was  rising  rather  than  being  forced  down- 
ward was  proved  by  its  regurgitation  immediately 
after  the  sound  was  heard.  In  all  probability  the 
pressure  that  forced  the  gas  from  the  stomach  was 
the  cause  of  the  preceding  sensation  of  hunger. 
Again  the  sensation  can  be  momentarily  abolished 
a  few  seconds  after  swallowing  a  small  accumula- 
tion of  saliva  or  a  teaspoonful  of  water.  If  the 
stomach  is  in  strong  contraction  in  hunger,  this  re- 
sult can  be  accounted  for,  in  accordance  with  the 
observations  of  Lieb  and  myself,^^  as  due  to  the 
inhibition  of  the  contraction  by  swallowing.  Thus 
also  could  be  explained  the  prompt  vanishing  of 
the  ache  soon  after  we  begin  to  eat,  for  repeated 
swallowing  results  in  continued  inhibition.*  Fur- 
thermore,   Ducceschi's    discovery  ^®    that    hydro- 

*  The  absence  of  hunger  in  Busch's  patient  while  food  was 
being  eaten  (see  p.  239)  can  also  be  accounted  for  in  this 
manner. 


THE    NATURE   OF   HUNGER         255 

chloric  acid  diminishes  the  tonus  of  the  pyloric  por- 
tion of  the  stomach  may  have  its  application  here ; 
the  acid  would  be  secreted  as  food  is  taken  and 
would  then  cause  relaxation  of  the  very  region 
which  is  most  strongly  contracted. 

The  Concomitance  of  Contractions  and  Hunger  in  Man 

Although  the  evidence  above  outlined  had  led 
me  to  the  conviction  that  hunger  results  from  con- 
tractions of  the  alimentary  canal,  direct  proof  was 
still  lacking.  In  order  to  learn  whether  such  proof 
might  be  secured,  Washburn  determined  to  be- 
come accustomed  to  the  presence  of  a  rubber  tube 
in  the  esophagus.*  Almost  every  day  for  several 
weeks  Washburn  introduced  as  far  as  the  stomach 
a  small  tube,  to  the  lower  end  of  which  was  attached 
a  soft-rubber  balloon  about  8  centimeters  in  diam- 
eter. The  tube  was  thus  carried  about  each  time 
for  two  or  three  hours.  After  this  preliminary^ 
experience  the  introduction  of  the  tube  and  its 
presence  in  the  gullet  and  stomach  were  not  at  all 
disturbing.  When  a  record  was  to  be  taken,  the 
l)al]oon,  placed  just  within  the  stomach,  was  moder- 
ately distended  with  air,  and  was  connected  with  a 
water  manometer  ending  in  a  cylindrical  chamber 
3.5  centimeters  wide.    A  float  recorder  resting  on 

*  Nicolai  (loc.  cit.)  reported  that  although  the  introduction 
of  a  stomach  tube  at  first  abolished  hunger  in  his  subjects, 
with  repeated  use  the  effects  became  insignificant. 


256  BODILY    CHANGES 

the  water  in  the  chamber  permitted  registering  any 
contractions  of  the  fundus  of  the  stomach.  On  the 
days  of  observation  Washburn  would  abstain  from 
breakfast,  or  eat  sparingly;  and  without  taking 
any  luncheon  would  appear  in  the  laboratory  about 
two  o'clock.  The  recording  apparatus  was  ar- 
ranged as  above  described.  In  order  to  avoid  any 
error  that  might  arise  from  artificial  pressure  on 
the  balloon,  a  pneumograph,  fastened  below  the 
ribs,  was  made  to  record  the  movements  of  the 
abdominal  wall.  Uniformity  of  these  movements 
would  show  that  no  special  contractions  of  the  ab- 
dominal muscles  were  made.  Between  the  records 
of  gastric  pressure  and  abdominal  movement,  time 
was  marked  in  minutes,  and  an  electromagnetic 
signal  traced  a  line  which  could  be  altered  by  press- 
ing a  key.  All  these  recording  arrangements  were 
out  of  Washburn's  sight ;  he  sat  with  one  hand  at 
the  key,  ready  whenever  the  sensation  of  hunger 
was  experienced  to  make  the  current  which  moved 
the  signal. 

Sometimes  the  observations  were  started  before 
any  hunger  was  noted;  at  other  times  the  sensa- 
tion, after  running  a  course,  gave  way  to  a  feeling 
of  fatigue.  Under  either  of  these  circumstances 
there  were  no  contractions  of  the  stomach.  When 
Washburn  stated  that  he  was  hungry,  however, 
powerful  contractions  of  the  stomach  were  invari- 
ably being  registered.    As  in  my  own  earlier  expe- 


THE   NATURE    OF   HUNGER         257 

rience,  the  sensations  were  characterized  by  peri- 
odic recurrences  with  free  intervals,  or  by  peri- 
odic accesses  of  an  uninterrupted  ache.  The  record 
of  Washburn's  introspection  of  his  hunger  pangs- 
agreed  closely  with  the  record  of  his  gastric  con- 


FiGURE  37.— One-half  the  original  size. 
The  top  record  represents  intragastric  pres- 
sure (the  small  oscillations  due  to  respiration, 
the  large  to  contractions  of  the  stomach);  the 
se(;ond  record  is  time  in  minutes  (ten  min- 
utes); the  third  record  is  W's  report  of  hunger 
pangs;  the  lowest  record  is  respiration  regis- 
tered by  means  of  a  pneumograph  about  the 
abdomen. 

tractions.  Almost  invariably,  however,  the  con- 
traction nearly  reached  its  maximum  before  the 
record  of  the  sensation  was  started  (see  Fig.  37). 
This  fact  may  be  regarded  as  evidence  that  the 
contraction  precedes  the  sensation,  and  not  vice 
versa,  as  Boldireff  considered  it.  The  contrac- 
tions were  about  a  half-minute  in  duration,  and 


258 


BODILY   CHANGES 


the  intervals  between  varied  from  thirty  to  ninety 
seconds,  with  an  average  of  about  one  minute.  The 
augmentations  of  intragastric  pressure  in  Wash- 
burn ranged  between  eleven  and  thirteen  in  twenty 
minutes ;  I  had  previously  counted  in  myself  eleven 
hunger  pangs  in  the  same  time.    The  rate  in  each 


Figure  38. — One-half  the  original  size.  The  same  conditions 
as  in  Fig.  37.  (Fifteen  minutes.)  There  was  a  long' wait  for  hunger 
to  disappear.  After  x,  W.  reported  himself  "tired  but  not  hungry." 
The  record  from  y  to  z  was  the  continuance,  on  a  second  drum,  of 
X  toy. 


of  US  was,  therefore,  approximately  the  same. 
This  rate  is  slightly  slower  than  that  found  in 
dogs  by  Boldireff ;  the  difference  is  perhaps  corre- 
lated with  the  slower  rhythm  of  gastric  peristalsis 
in  man  compared  with  that  in  the  dog.^''' 

Before  hunger  was  experienced  by  Washburn 
the  recording  apparatus  revealed  no  signs  of  gas- 
tric activity.  Sometimes  a  rather  tedious  period 
of  waiting  had  to  be  endured  before  contractions 


THE    NATURE    OF   HUNGER         259 

occurred.  And  after  tliey  began  they  continued 
for  a  while,  then  ceased  (see  Fig.  38).  The  feeling 
of  hunger,  which  was  reported  while  the  contrac- 
tions were  recurring,  disappeared  as  the  waves 
stopped.  The  inability  of  the  subject  to  control 
the  contractions  eliminated  the  possibility  of  their 
being  artifacts,  perhaps  induced  by  suggestion. 
The  close  concomitance  of  the  contractions  with 
hunger  pangs,  therefore,  clearly  indicates  that  they 
are  the  real  source  of  those  pangs. 

Boldiretf's  studies  proved  that  when  the  empty 
stomach  is  manifesting  periodic  contractions,  the 
intestines  also  are  active.  Conceivably  all  parts 
of  the  alimentary  canal  composed  of  smooth  mus- 
cle share  in  these  movements.  The  lower  esopha- 
gus in  man  is  provided  with  smooth  muscle.  It 
was  possible  to  determine  whether  this  region  in 
Washburn  was  active  during  hunger. 

To  the  esophageal  tube  a  thin-rubber  finger-cot 
(2  centimeters  in  length)  was  attached  and  lowered 
into  the  stomach.  The  little  rubber  bag  was  dis- 
tended with  air,  and  the  tube,  pinched  to  keep  the 
bag  inflated,  was  gently  withdra^\Ti  until  resistance 
was  felt.  The  air  was  now  released  from  the  bag 
and  the  tube  farther  withdrawn  about  3  centi- 
meters. The  l)ag  was  again  distended  with  air  at 
a  manometric  pressure  of  10  centimeters  of  water. 
Inspiration  now  caused  the  writing  lever,  which 
recorded   the  pressure  changes,  to   rise;   and  a 


260  BODILY   CHANGES 

slightly  farther  withdrawal  of  the  tube  changed 
the  rise,  on  inspiration,  to  a  fall.  The  former  posi- 
tion of  the  tnbe,  therefore,  was  above  the  gastric 
cavity  and  below  the  diaphragm.    In  this  position 


Figure  39. — One-half  the  original  size. 
The  top  record  represents  compression  of 
thin  rubber  bag  in  the  lower  esophagus. 
The  pressure  in  the  bag  varied  between  9 
and  13  centimeters  of  water.  The  cyUn- 
der  of  the  recorder  was  of  smaller  diameter 
than  that  used  in  the  gastric  records.  The 
esophageal  contractions  compressed  the 
bag  so  completely  that,  at  the  summits  of 
the  large  oscillations,  the  respirations  were 
not  registered.  When  the  oscillations 
dropped  to  the  time  hne,  the  bag  was  about 
half  inflated.  The  middle  Une  registers 
time  in  minutes  (ten  minutes).  The  bot- 
tom record  is  W's  report  of  hunger  pangs. 


the  bag,  attached  to  a  float  recorder  (with  chamber 
2.3  centimeters  in  diameter),  registered  the  peri- 
odic oscillations  shown  in  Fig.  39.  Though  indi- 
vidually more  prolonged  than  those  of  the  stomach, 
these  contractions,  it  will  be  noted,  occur  at  about 
the  same  rate. 


THE   NATURE    OF   HUNGER         261 

This  study  of  hunger,  reported  by  Washburn 
and  myself  in  1912,  has  since  been  taken  up  by 
Carlson  of  Chicago,  and  in  observations  on  a  man 
with  a  permanent  gastric  fistula,  as  well  as  on  him- 
self and  his  collaborators,  he  has  fully  confinned 
our  evidence  as  to  the  relation  between  contrac- 
tions of  the  alimentary  canal  and  the  hunger  sensa- 
tion. In  a  series  of  nearly  a  score  of  interesting 
papers,  Carlson  and  his  students  ^^  have  greatly 
amplified  our  knowledge  of  the  physiology  of  the 
"empty"  stomach.  Not  only  are  there  the  contrac- 
tions observed  by  Washburn  and  myself,  but  at 
times  these  may  fuse  into  a  continuous  cramp  of 
the  gastric  muscle.  The  characteristic  contrac- 
tions, furthermore,  continue  after  the  vagus  nerve 
supply  to  the  stomach  has  been  destroyed,  and, 
therefore,  are  not  dependent  on  the  reception  of 
impulses  by  way  of  the  cranial  autonomic  fibres. 
Recently  Luckhardt  and  Carlson  have  brought  for- 
ward evidence  that  the  blood  of  a  fasting  animal 
if  injected  into  the  vein  of  a  normal  animal  is 
capable  of  inducing  in  the  latter  the  condition  of 
cramp  or  tetanus  in  the  gastric  muscle  mentioned 
above — an  effect  which  does  not  occur  when  the 
blood  of  a  well-fed  animal  is  injected.  It  seems 
possible  that  a  substance  exists  in  the  blood  which 
acts  to  excite  the  gastric  hunger  mechanism.  But 
this  point  will  require  further  investigation. 

With  these  demonstrations  that  contractions  are 


262  BODILY   CHANGES 

the  immediate  cause  of  hniiger,  most  of  the  diffi- 
culties confronting  other  explanations  are  readily 
obviated.  Thus  the  sudden  onset  of  hunger  and 
its  peculiar  periodicity— phenomena  which  no 
other  explanation  of  hunger  can  account  for— are 
at  once  explained. 

In  fever,  when  bodily  material  is  being  most 
rapidly  used,  hunger  is  absent.  Its  absence  is 
understood  from  an  observation  made  by  F.  T. 
Murphy  and  myself,^*^  that  infection,  with  sys- 
temic involvement,  is  accompanied  by  a  total 
cessation  of  all  movements  of  the  alimentary  canal. 
Boldireff  observed  that  when  his  dogs  were  fa- 
tigued the  rhythmic  contractions  failed  to  appear. 
Being  "too  tired  to  eat"  is  thereby  given  a  rational 
explanation. 

A  pathological  form  of  the  sensation — the  inor- 
dinate hunger  (bulimia)  of  certain  neurotics — is 
in  accordance  with  the  well-known  disturbances 
of  the  tonic  innervation  of  the  alimentary  canal 
in  such  individuals. 

Since  the  lower  end  of  the  esophagus,  as  well 
as  the  stomach,  contracts  periodically  in  hunger, 
the  reference  of  the  sensation  to  the  sternum  by 
the  ignorant  persons  questioned  by  Schitf  was 
wholly  natural.  The  activity  of  the  lower  esopha- 
gus also  explains  why,  after  the  stomach  has  been 
removed,  or  in  some  cases  when  the  stomach  is 
distended  with  food,  hunger  can  still  be  experi- 


THE   NATUIiE    OF   HUNGER         263 

enced.  Conceivably  the  intestines  also  originate 
vague  sensations  by  their  contractions.  Indeed, 
the  final  banishment  of  the  modified  hunger  sen- 
sation in  the  patient  with  duodenal  fistula,  de- 
scribed by  Busch,  may  have  been  due  to  the  les- 
sened activity  of  the  intestines  when  chyme  was 
injected  into  them. 

The  observations  recorded  in  this  paper  have, 
as  already  noted,  numerous  points  of  similarity  to 
Boldireff's  observations  ^"  on  the  periodic  activ- 
ity of  the  alimentary  canal  in  fasting  dogs.  Each 
period  of  activity,  he  found,  comprised  not  only 
wide-spread  contractions  of  the  digestive  canal,  but 
also  the  pouring  out  of  bile,  and  of  pancreatic  and 
intestinal  juices  rich  in  ferments.  Gastric  juice 
vras  not  secreted  at  these  times;  when  it  was  se- 
creted and  reached  the  intestine,  the  periodic  activ- 
ity ceased.  Wliat  is  the  significance  of  this  exten- 
sive disturbance  I  I  have  elsewhere  presented  evi- 
dence ^^  that  gastric  peristalsis  is  dependent  on 
the  stretching  of  gastric  muscle  when  tonically  con- 
tracted. The  evidence  that  the  stomach  is  in  fact 
strongly  contracted  in  hunger — i.  e.,  in  a  state  of 
high  tonus — has  been  presented   above.*     Thus 

*  The  "empty"  stomach  and  esophagus  contain  gas  (see 
Hertz:  Quarterly  Journal  of  Medicine,  1910,  iii,  p.  378; 
Mikulicz:  Mittheilungen  aus  den  Grenzgebieten  der  Medi- 
cin  und  Chirurgie,  1903,  xii,  p.  590).  They  would  naturally 
manifest  rhythmic  contractions  on  shortening  tonically  on 
their  content. 


264  BODILY   CHANGES 

the  very  condition  which  causes  hunger  and  leads 
to  the  taking  of  food  is  the  condition,  when  the 
swallowed  ^ood  stretches  the  shortened  muscles, 
for  immediate  starting  of  gastric  peristalsis.  In 
this  connection  the  observations  of  Haudek  and 
Stigler*^  are  probably  significant.  They  found 
that  the  stomach  discharges  its  contents  more  rap- 
idly if  food  is  eaten  in  hunger  than  if  not  so  eaten. 
Hunger,  in  other  words,  is  normally  the  signal  that 
the  stomach  is  contracted  for  action ;  the  unpleas- 
antness of  hunger  leads  to  eating;  eating  starts 
gastric  digestion,  and  abolishes  the  sensation. 
Meanwhile  the  pancreatic  and  intestinal  juices,  as 
well  as  bile,  have  been  prepared  in  the  duodenum 
to  receive  the  oncoming  chyme.  The  periodic  ac- 
tivity of  the  alimentary  canal  in  fasting,  therefore, 
is  not  solely  the  source  of  hunger  pangs,  but  is  at 
the  same  time  an  exhibition  in  the  digestive  organs 
of  readiness  for  prompt  attack  on  the  food  swal- 
lowed by  the  hungry  animal. 

EEFERENCES 

^  Cannon :  The  Mechanical  Factors  of  Digestion,  London 
and  New  York,  1911,  p.  204. 

2  Bardier :  Eichet's  Dictionnaire  de  Physiologie,  article 
Eaim,  1904,  vi,  p.  1.  See,  also,  Howell :  Text-book  of  Physi- 
ology, fourth  edition,  Philadelphia  and  London,  1911,  p.  285. 

^  See  Sternberg :  Zentralblatt  f iir  Physiologie,  1909, 
xxii,  p.  653.  Similar  views  were  expressed  by  Bayle  in  a 
thesis  presented  to  the  Faculty  of  Medicine  in  Paris  in  1816. 

*  See  Hertz :  The  Sensibility  of  the  Alimentary  Canal, 
London,  1911,  p.  38. 


THE   NATURE    OF   HUNGER         265 

^  SchiflF :  Physiologie  de  la  Digestion,  Florence  and  Turin, 
1867,  p.  40. 

®  Luciani :  Das  Hungem,  Hamburg  and  Leipzig,  1890, 
p.  113. 

"^  Tigerstedt :  Nagel's  Handbueh  der  Physiologie,  Berlin, 
1909,  i,  p.  376. 

*  Johanson,  Landergreu,  Sonden  and  Tigerstedt :  Skandi- 
navisches  Archiv  fiir  Physiologie,  1897,  vii,  p.  33. 

^  Carrington :  Vitality,  Fasting  and  Nutrition,  New 
York,  1908,  p.  555. 

^°  Viterbi,  quoted  by  Bardier :  Loc.  cit.,  p.  7. 

^^  Busch :  Archiv  fiir  pathologische  Anatomie  und  Physi- 
ologie und  fiir  klinische   Medicin,   1858,  xiv,  p.   147. 

^^  See  Schiff:  Loc.  cit.,  p.  37;  also  Ducceschi;  Archivio  di 
Fisiologia,  1910,  viii,  p.  579. 

^^  Longet :  Traite  de  Physiologie,  Paris,  1868,  i,  p.  23. 

^*  Ludwig :  Lehrbuch  der  Physiologie  des  Menschen,  Leip- 
zig and  Heidelberg,   1858,  ii,  p.   584. 

^^  Maxwell :  Journal  of  Biological  Chemistry,  1906-7,  ii, 
p.  194. 

16  See    SchiflF:   Loc.    cit.,   p.   49. 

i'^  See  SchiflF:  Loc.  cit.,  p.  31;  Bardier;  Loc.  cit.,  p.  16. 

18  Head:  Brain,  1893,  xvi,  p.  1;  1901,  xxiv,  p.  345. 

1^  Nicolai :  Ueber  die  Entstehung  des  Hungergefiihls,  In- 
augural Dissertation,  Berlin,  1892,  p.  17. 

**^  Beaumont :  The  Physiology  of  Digestion,  second  edi- 
tion, Burlington,  1847,  p.  51. 

21  Nicolai :  Loc.  cit.,  p.  15. 

22  Beaumont:  Loc.  cit.,  p.  55. 

23  Luciani :  Archivio  di  Fisiologia,  1906,  iii,  p.  54.  Tiede- 
mann  long  ago  suggested  that  gastric  nerves  become  increas- 
ingly sensitive  as  fasting  progresses.  (Physiologie  des  Men- 
schen, Darmstadt,  1836,  iii,  p.  22.) 

2*  Valenti :  Archives  Italiennes  de  Biologie,  1910,  liii,  p.  94. 
2^  Weber :  Wagner's  Handworterbuch  der  Physiologie,  1846, 
iii2,  p.  580. 

26  Vierordt :  Grundriss  der  Physiologie,  Tiibingen,  1871, 
p.  433. 

27  SchiflF :  Loc.  cit.,  p.  33. 


266  BODILY    CHANGES 

28Luciani:  Loc.  cit.,  p.  542. 

29  Valenti :  Loc.  cit.,  p.  95. 

30  Bettmann :  Philadelphia  Monthly  Medical  Journal,  1899, 
i,  p.  133. 

31  Wolff:  Dissertation,  Giessen,  1902,  p.  9. 

32  His :  Archiv  fur  Anatomie,  1903,  p.  345. 

33  Boldireff :  Loc.  cit.,  p.  1. 
3*  Boldireff :  Loc.  cit.,  p.  96. 

3^  See  Cannon  and  Lieb :  American  Journal  of  Physiol- 
ogy, 1911,  xxix,  p.  267. 

3''  Ducceschi :  Archivio  per  le  Scienze  Mediche,  19Y,  xxi, 
p.  154. 

3'''  See  Cannon :  American  Journal  of  Physiology,  1903, 
viii,  p.  xxi ;  1905,  xiv,  p.  344. 

3^  See  American  Journal  of  Physiology,  1913,  1914. 

39  Cannon  and  Murphy :  Journal  of  the  American  Medi- 
cal Association,  1907,  xlix,  p.  840. 

*"  Boldireff :  Loc.  cit.,  pp.  108-111. 

^1  Cannon :  American  Journal  of  Physiology,  1911,  xxix, 
p.  250. 

*^  Haudek  and  Stigler :  Archiv  f  iir  die  gesammte  Physi- 
ologic, 1910,  cxxxiii,  p.  159. 


CHAPTER   XIV 

THE   INTERKELATIONS   OF   EMOTIONS 

Emotions  gain  expression  through  discharges 
along  the  neurones  of  the  autonomic  nervous  sys- 
tem. The  reader  will  recall  that  this  system  has 
three  divisions — the  cranial  and  sacral,  separated 
by  the  sympathetic — and  that  when  the  neurones  of 
the  mid-division  meet  in  any  organ  the  neurones  of 
either  of  the  end  divisions,  the  influence  of  the  two 
sets  is  antagonistic.  As  previously  stated  (p.  35), 
there  is  evidence  that  arrangements  exist  in  the 
central  nerv^ous  system  for  reciprocal  innervation 
of  these  antagonistic  divisions,  just  as  there  is 
reciprocal  innervation  of  antagonistic  skeletal 
muscles.  The  characteristic  atfective  states  mani- 
fested in  the  working  of  these  three  divisions  have 
been  described.  Undoubtedly,  these  states  have 
correspondents — activities  and  inhibitions — in  the 
central  neurones.  The  question  now  arises,  are 
the  states  which  appear  in  opposed  divisions  also 
in  opposition? 

267 


268  BODILY    CHANGES 

Antagonism  Between  Emotions  Expressed  in  the  Sympa- 
thetic AND  IN  the  Cranial  Divisions  op  the  Autonomic 
System 

The  cranial  autonomic,  as  already  shown,  is  con- 
cerned with  the  quiet  service  of  building  up  re- 
serves and  fortifying  the  body  against  times  of 
stress.  Accompanying  these  functions  are  the 
relatively  mild  pleasures  of  sight  and  taste  and 
smell  of  food.  The  possibility  of  existence  of  these 
gentle  delights  of  eating  and  drinking  and  also  of 
their  physiological  consequences  is  instantly  abol- 
ished in  the  presence  of  emotions  which  activate 
the  sympathetic  division.  The  secretion  of  saliva, 
gastric  juice,  pancreatic  juice  and  bile  is  stopped, 
and  the  motions  of  the  stomach  and  intestines  cease 
at  once,  both  in  man  and  in  the  lower  animals, 
whenever  pain,  fear,  rage,  or  other  strong  excite- 
ment is  present  in  the  organism. 

All  these  disturbances  of  digestion  seem  mere 
interruptions  of  the  "normal"  course  of  events 
unless  the  part  they  may  play  in  adaptive  reac- 
tions is  considered.  In  discussing  the  operations 
of  the  sympathetic  division,  I  pointed  out  that  all 
the  bodily  changes  which  occur  in  the  intense  emo- 
tional states — such  as  fear  and  fury — occur  as 
results  of  activity  in  this  division,  and  are  in  the 
highest  degree  serviceable  in  the  struggle  for  exist- 
ence likely  to  be  precipitated  when  these  emotions 
are  aroused.    From  this  point  of  view  these  per- 


INTERRELATIONS  OF  EMOTIONS    269 

turbations,  which  so  readily  seize  and  dominate 
the  organs  that  in  quiet  times  are  commonly  con- 
trolled by  the  cranial  autonomic,  are  bodily  reac- 
tions which  may  be  of  the  utmost  importance  to  life 
at  times  of  critical  emergency.  Thus  are  the  body's 
reserves — the  stored  adrenin  and  the  accumulated 
sugar — called  forth  for  instant  service ;  thus  is  the 
blood  shifted  to  nerves  and  muscles  that  may  have 
to  bear  the  brunt  of  strugg-le ;  thus  is  the  heart  set 
rapidly  beating  to  speed  the  circulation ;  and  thus, 
also,  are  the  activities  of  the  digestive  organs  for 
the  time  abolished.  Just  as  in  war  between  nations 
the  arts  and  industries  which  have  brought  wealth 
and  contentment  must  suffer  serious  neglect  or  be 
wholly  set  aside  both  by  the  attacker  and  the  at- 
tacked, and  all  the  supplies  and  energies  developed 
in  the  period  of  peace  must  be  devoted  to  the  pres- 
ent conflict;  so,  likewise,  the  functions  which  in 
quiet  times  establish  and  support  the  bodily  re- 
serves are,  in  times  of  stress,  instantly  checked 
or  completely  stopped,  and  these  reserves  lavishly 
drawn  upon  to  increase  power  in  the  attack  and  in 
the  defense  or  flight.* 

It  is,  therefore,  the  natural  antagonism  between 
these  two  processes  in  the  body — between  saving 

*  One  who  permits  fears,  worries  and  anxieties  to  disturb 
the  digestive  processes  when  there  is  nothing  to  be  done,  is 
evidently  allowing  the  body  to  go  onto  what  we  may  regard 
as  a  "war  footing,"  when  there  is  no  "war"  to  be  waged,  no 
fighting  or  struggle  to  be  engaged  in. 


270  BODILY   CHANGES 

and  expenditure,  between  preparation  and  use,  be- 
tween anabolism  and  catabolism — and  the  corre- 
lated antagonism  of  central  innervations,  that  un- 
derlie the  antipathy  between  the  emotional  states 
which  normally  accompany  the  processes.  The 
desire  for  food,  the  relish  of  eating  it,  all  the 
pleasures  of  the  table,  are  naught  in  the  presence 
of  anger  or  great  anxiety.  And  of  the  two  sorts 
of  emotional  states,  those  which  manifest  them- 
selves in  the  dominant  division  of  the  autonomic 
hold  the  field  also  in  consciousness. 

Antagonism  Between  Emotions  Expressed  in  the  Sympa- 
thetic AND  IN   THE   SaCRAL  DIVISIONS  OP  THE  AUTONOMIC 

System 

The  nervi  erigentes  are  the  part  of  the  sacral 
autonomic  in  which  the  peculiar  excitements  of 
sex  are  expressed.  As  previously  stated,  these 
nerves  are  opposed  by  branches  from  the  sympa- 
thetic division — the  division  which  is  operated 
characteristically  in  the  major  emotions. 

The  opposition  in  normal  individuals  between  the 
emotional  states  which  appear  in  these  two  antag- 
onistic divisions  is  most  striking.  Even  in  animals 
as  low  in  the  scale  as  birds,  copulation  is  not  per- 
formed "until  every  condition  of  circumstance  and 
sentiment  is  fulfilled,  until  time,  place  and  partner 
all  are  fit."^  And  among  men  the  effect  of  fear 
or  momentary  anxiety  or  any  intense  emotional 
interest  in  causing  inhibition  of  the  act  can  be  sup- 


INTERKELATIONS  OF  EMOTIONS    271 

ported  by  cases  in  the  experience  of  any  physician 
with  extensive  practice.  Indeed,  as  Prince  -  has 
stated,  "the  suppression  of  the  sexual  instinct  by 
conflict  is  one  of  the  most  notorious  experiences 
of  this  kind  in  everyday  life.  This  instinct  cannot 
be  excited  during  an  attack  of  fear  or  anger,  and 
even  during  moments  of  its  excitation,  if  there  is 
an  invasion  of  another  strong  emotion  the  sexual 
instinct  at  once  is  repressed.  Under  these  con- 
ditions, as  with  other  instincts,  even  habitual 
excitants  can  no  longer  initiate  the  instinctive 
process." 

When  the  acme  of  excitement  is  approaching  it 
is  probable  that  the  sympathetic  division  is  also 
called  into  activity ;  indeed,  the  completion  of  the 
process — the  contractions  of  the  seminal  vesicles 
and  the  prostate,  and  the  subsidence  of  engorged 
tissues,  all  innervated  by  sympathetic  filaments 
(see  pp.  32,  33) — may  be  due  to  the  overwhelm- 
ing of  sacral  by  sympathetic  nervous  discharges. 
As  soon  as  this  stage  is  reached  the  original  feeling 
likewise  has  been  dissipated. 

The  other  parts  of  the  sacral  division  which 
supply  the  bladder  and  rectum  are  so  nearly  free 
from  any  emotional  tone  in  their  normal  reflex 
functioning  that  it  is  unnecessary  to  consider  them 
further  with  reference  to  emotional  antagonisms. 
Mild  affective  states,  such  as  worry  and  anxiety, 
can,  to  be  sure,  check  the  activity  of  the  colon  and 


272  BODILY   CHANGES 

thus  cause  constipation.^  But  the  augumented 
activity  of  these  parts  (contraction  of  the  bladder 
and  rectum)  in  very  intense  periods  of  emotional 
stress,  when  the  sympathetic  division  is  strongly 
innervated,  presents  a  problem  of  some  difficulty. 
Possibly  in  such  conditions  the  orderliness  of  the 
central  arrangements  is  upset,  just  as  it  is  after 
tetanus  toxin  or  strychnine  poisoning,  and  opposed 
innervations  no  longer  discharge  reciprocally,  but 
simultaneously,  and  then  the  stronger  member  of 
the  pair  prevails.  Only  on  such  a  basis,  at  pres- 
ent, can  I  offer  any  explanation  for  the  activity 
and  the  supremacy  of  the  sacral  innervation  of 
the  bladder  and  distal  colon  when  the  sympathetic 
innervation  is  aroused,  as,  for  example,  in  great 
fright. 

The  Function  of  Hunger 

A  summary  in  few  words  of  the  chief  functions 
typically  performed  or  supported  by  each  division 
of  the  autonomic  would  designate  the  cranial  divi- 
sion as  the  upbuilder  and  restorer  of  the  organic 
reserves,  the  sacral  as  the  servant  of  racial  contin- 
uity, and  the  sympathetic  as  the  preserver  of  the  in- 
dividual. Self-preservation  is  primary  and  essen- 
tial ;  on  that  depends  racial  continuity,  and  for  that 
all  the  resources  of  the  organism  are  called  forth. 
Analogously  the  sympathetic  innervations,  when 
they  meet  in  organs  innervated  also  by  the  cranial 


INTERRELATIONS  OF  EMOTIONS    273 

and  sacral  divisions,  almost  without  exception  pre- 
dominate over  their  opponents.  And  analogously, 
also,  the  emotional  states  which  are  manifested  in 
the  sympathetic  division  and  are  characteristically 
much  more  intense  than  those  manifested  in  the 
other  divisions,  readily  assume  ascendancy  also  in 
consciousness. 

It  is  obvious  that  extended  action  of  the  sym- 
pathetic division,  abolishing  those  influences  of  the 
cranial  division  which  are  favorable  to  proper  di- 
gestion and  nutrition,  might  defeat  its  own  ends. 
Interruption  of  the  nutritional  process  for  the  sake 
of  self-preservation  through  defense  or  attack  can 
l)e  only  temporary;  if  the  interruption  were  pro- 
longed, there  might  be  serious  danger  to  the  vigor 
of  the  organism  from  failure  to  replenish  the  ex- 
hausted stores.  The  body  does  not  have  to  depend 
on  the  return  of  a  banished  appetite,  however,  be- 
fore its  need  for  restoration  is  attended  to.  There 
is  a  secondary  and  very  insistent  manner  in  which 
the  requirement  of  food  is  expressed,  and  that  is 
through  the  repeated  demands  of  hunger. 

Unlike  many  other  rhythmically  repeated  sen- 
sations, hunger  is  not  one  that  anybody  becomes  ac- 
customed to  and  neglects  because  of  its  monotony. 
During  the  period  of  his  confinement  in  the  citadel 
of  Magdeburg,  the  celebrated  jjolitical  adventurer 
Baron  von  Trenck  '*  was  allowed  only  a  pound 
and  a  half  of  ammunition  bread  and  a  jug  of  water 


274  BODILY   CHANGES 

as  his  daily  ration.  "It  is  impossible  for  me  to 
describe  to  my  reader,"  he  wrote  in  his  memoirs, 
"the  excess  of  tortures  that  during  eleven  months 
I  endured  from  ravenous  hunger.  I  could  easily 
have  devoured  six  pounds  of  bread  every  day ;  and 
every  twenty-four  hours,  after  having  received 
and  swallowed  my  small  portion,  I  continued  as 
hungry  as  before  I  began,  yet  I  was  obliged  to  wait 
another  twenty-four  hours  for  a  new  morsel. 
.  .  .  My  tortures  prevented  sleep,  and  looking 
into  futurity,  the  cruelty  of  my  fate  seemed  to  me, 
if  possible,  to  increase,  for  I  imagined  that  the  pro- 
longation of  pangs  like  these  was  insupportable. 
God  preserve  every  honest  man  from  sufferings 
like  mine!  They  were  not  to  be  endured  by  the 
most  obdurate  villain.  Many  have  fasted  three 
days,  many  have  suffered  want  for  a  week  or  more, 
but  certainly  no  one  besides  myself  ever  endured 
it  in  the  same  excess  for  eleven  months ;  some  have 
supposed  that  to  eat  little  might  become  habitual, 
but  I  have  experienced  the  contrary.  My  hunger 
increased  every  day,  and  of  all  the  trials  of  forti- 
tude my  whole  life  has  afforded,  this  eleven  months 
was  the  most  bitter."* 

*  In  all  probability  the  continued  experience  of  hunger 
pangs  reported  by  Baron  von  Trenck  was  due  to  the  re- 
peated eating  of  amounts  of  food  too  small  to  satisfy  the 
bodily  demand.  The  reader  will  recall  that  persons  who  for 
some  time  take  no  food  whatever  report  that  the  disagreeable 
feelings  are  less  intense  or  disappear  after  the  third  or  fourth 
day   (see  p.   238). 


INTERRELATIONS  OF  EMOTIONS    275 

Thus,  although  the  taking  of  food  may  be  set  in 
abeyance  at  times  of  great  excitement,  and  the 
bodily  reserves  fully  mobilized,  that  phase  of  the 
organism's  self-protecting  adjustment  is  limited, 
and  then  hunger  asserts  itself  as  an  agency  im- 
periously demanding  restoration  of  the  depleted 
stores. 

The  Similarity  of  Visceral  Effects  in  Different  Stronq 

Emotions  and  Suggestions  as  to  its  Psychological 

Significance 

The  dominant  emotions  which  we  have  been  con- 
sidering as  characteristically  expressed  in  the 
sjTnpathetic  division  of  the  autonomic  system  are 
fear  and  rage.  These  two  emotions  a're  not  unlike. 
As  James  ^  has  indicated,  "Fear  is  a  reaction 
aroused  by  the  same  objects  that  arouse  ferocity. 
.  .  .  We  both  fear  and  wish  to  kill  anything 
that  may  kill  us ;  and  the  question' which  of  the  two 
impulses  we  shall  follow  is  usually  decided  by  some 
one  of  those  collateral  circumstances  of  the  par- 
ticular case,  to  be  moved  by  which  is  the  mark  of 
superior  mental  natures."  The  cornering  of  an 
animal  when  in  the  headlong  flight  of  fear  may 
suddenly  turn  the  fear  to  fury  and  the  flight  to  a 
fighting  in  which  all  the  strength  of  desperation 
is  displayed. 

Furthermore,  these  dominant  emotions  are  states 
into  which  many  other  commonly  milder  affective 
states  may  be  suddenly  transformed.    As  McDou- 


276  BODILY    CHANGES 

gall^  lias  pointed  out,  all  instinctive  impulses 
when  met  with  opposition  or  obstruction  give  place 
to,  or  are  complicated  by,  the  pugnacious  or  com- 
bative impulse  directed  against  the  source  of  the 
obstruction.  A  dog  will  bristle  at  any  attempt  to 
take  away  his  food,  males  will  fight  furiously  when 
provoked  by  interference  with  the  satisfaction  of 
the  sexual  impulse,  a  man  will  forget  the  conven- 
tions and  turn  hot  for  combat  when  there  is  impu- 
tation against  his  honor,  and  a  mother  all  gentle 
with  maternal  devotion  is  stung  to  quick  resent- 
ment and  will  make  a  fierce  display  of  her  com- 
bative resources,  if  anyone  intentionally  injures 
her  child.  In  these  instances  of  thwarted  or  dis- 
turbed instinctive  acts  the  emotional  accompani- 
ments— such  as  the  satisfaction  of  food  and  of 
sexual  affection,  the  feeling  of  self -pride,  and  the 
tender  love  of  a  parent — are  whirled  suddenly  into 
anger.  And  anger  in  one  is  likely  to  provoke 
anger  or  fear  in  the  other  who  for  the  moment  is 
the  object  of  the  strong  feeling  of  antagonism. 
Anger  is  the  emotion  preeminently  serviceable 
for  the  display  of  power,  and  fear  is  often  its 
/counterpart. 

The  visceral  changes  which  accompany  fear  and 
rage  are  the  result  of  discharges  by  way  of  sym- 
pathetic neurones.  It  will  be  recalled  that  these 
neurones  are  arranged  for  diffuse  rather  than  for 
narrowly  directed  effects.     So  far  as  these  two 


INTERRELATIONS  OF  EMOTIONS    277 

quite  different  emotions  are  concerned,  present 
physiological  evidence  indicates  that  differences 
in  visceral  accompaniments*  are  not  noteworthy — 
for  example,  either  fear  or  rage  stops  gastric  se- 
cretion (see  pp.  10,  11).  There  is,  indeed,  obvious 
reason  why  the  visceral  changes  in  fear  and  rage 
should  not  be  different,  but  rather,  why  they  should 
be  alike.  As  already  pointed  out,  these  emotions 
accompany  organic  preparations  for  action,  and 
just  because  the  conditions  which  evoke  them  are 
likely  to  result  in  flight  or  conflict  (either  one 
requiring  perhaps  the  utmost  struggle),  the  bodily 
needs  in  either  response  are  precisely  the  same. 

In  discussing  the  functioning  of  the  sympathetic 
division  I  jwinted  out  that  it  was  roused  to  ac- 
tivity not  only  in  fear  and  rage,  but  also  in  pain. 
The  machinery  of  this  division  likewise  is  oper- 
ated wholly  or  partially  in  emotions  which  are 
usually  mild — such  as  joy  and  sorrow  and  disgust 

•Obvious  vascular  differences,  as  pallor  or  flushing  of  the 
face,  are  of  little  significance.  With  increase  of  blood  pres- 
sure from  vasoconstriction,  pallor  might  result  from  action  of 
the  constrictors  in  the  face,  or  flushing  might  result  because 
constrictors  elsewhere,  as,  for  example,  in  the  abdomen,  raised 
the  pressure  so  high  that  facial  constrictors  are  overcome. 
Such,  apparently,  is  the  effect  of  adrenin  already  described 
(see  p.  107).  Or  the  flushing  might  occur  from  local  vasodila- 
tion. That  very  different  emotional  states  may  hav'C  the  same 
vascular  accompaniments  was  noted  by  Darwin  (The  Expres- 
sion of  Emotions  in  Man  and  Animals,  New  York,  1905), 
who  mentioned  the  pallor  of  rage  (p.  74)  and  also  of  terror 
(p.  77). 


278  BODILY    CHANGES 

— when  they  become  sufftciently  intense.  Thus, 
for  instance,  the  normal  course  of  digestion  may 
be  stopped  or  quite  reversed  in  a  variety  of  these 
emotional  states. 

Darwin  '^  reports  the  case  of  a  young  man  who 
on  hearing  that  a  fortune  had  just  been  left  him, 
became  pale,  then  exhilarated,  and  after  various 
expressions  of  joyous  feeling  vomited  the  half- 
digested  contents  of  his  stomach.  Miiller^  has 
described  the  case  of  a  young  woman  whose  lover 
had  broken  the  engagement  of  marriage.  She 
wept  in  bitter  sorrow  for  several  days,  and  during 
that  time  vomited  whatever  food  she  took.  And 
Burton,''  in  his  Anatomy  of  Melancholy,  gives 
the  following  instance  of  the  effect  of  disgust :  "A 
gentlewoman  of  the  same  city  saw  a  fat  hog  cut 
up,  when  the  entrails  were  opened,  and  a  noisome 
savour  offended  her  nose,  she  much  misliked,  and 
would  not  longer  abide;  a  physician  in  presence 
told  her,  as  that  hog,  so  was  she,  full  of  filthy  ex- 
crements, and  aggravated  the  matter  by  some 
other  loathsome  instances,  insomuch  this  nice  gen- 
tlewoman apprehended  it  so  deeply  that  she  fell 
forthwith  a  vomiting,  was  so  mightily  distempered 
in  mind  and  body,  that  with  all  his  art  and  persua- 
sion, for  some  months  after,  he  could  not  restore 
her  to  herself  again,  she  could  not  forget  or  remove 
the  object  out  of  her  sight." 

In  these  three  cases,  of  intense  joy,  intense  sor- 


INTERRELATIONS  OF  EMOTIONS    279 

row  and  intense  disgust,  the  influence  of  tlie  cran- 
ial division  of  the  autonomic  has  been  overcome, 
digestion  has  ceased,  and  the  stagnant  gastric 
contents  by  reflexes  in  striated  muscles  have  been 
violently  discharged.  The  extent  to  which  under 
such  circumstances  other  effects  of  sympathetic 
impulses  may  be  manifested,  has  not,  so  far  as  I 
know,  been  ascertained. 

From  the  evidence  just  given  it  appears  that 
any  high  degree  of  excitement  in  the  central  nerv- 
ous system,  whether  felt  as  anger,  terror,  pain, 
anxiety,  joy,  grief  or  deep  disgust,  is  likely  to 
break  over  the  threshold  of  the  sympathetic  divi- 
sion and  disturb  the  functions  of  all  the  organs 
which  that  division  innervates.  It  may  be  that 
there  is  advantage  in  the  readiness  with  which 
these  widely  different  emotional  conditions  can  ex- 
press themselves  in  this  one  division,  for,  as  has 
been  shown  (see  p.  276),  occasions  may  arise  when 
these  milder  emotions  are  suddenly  transmuted 
into  the  naturally  intense  types  (as  fright  and 
fury)  which  normally  activate  this  division;  and 
if  the  less  intense  can  also  influence  it,  the  physio- 
logical aspect  of  the  transmutation  is  already  par- 
tially accomplished. 

If  various  strong  emotions  can  thus  be  expressed 
in  the  diffused  activities  of  a  single  division  of  the 
autonomic  —  the  division  which  accelerates  the 
heart,  inhibits  the  movements  of  the  stomach  and 


280  BODILY   CHANGES 

intestines,  contracts  the  blood  vessels,  erects  the 
hairs,  liberates  sugar,  and  discharges  adrenin — it 
would  appear  that  the  bodily  conditions  which  have 
been  assumed,  by  some  psychologists,  to  distin- 
guish emotions  from  one  another  must  be  sought 
for  elsewhere  than  in  the  viscera.  We  do  not  "feel 
sorry  because  we  cry,"  as  James  contended,  but 
we  cry  because  when  we  are  sorry  or  overjoyed  or 
violently  angry  or  full  of  tender  affection — ^when 
any  one  of  these  diverse  emotional  states  is  present 
— there  are  nervous  discharges  by  sympathetic 
channels  to  various  viscera,  including  the  lachry- 
mal glands.  In  terror  and  rage  and  intense  elation, 
for  example,  the  responses  in  the  viscera  seem  too 
uniform  to  offer  a  satisfactory  means  of  distin- 
guishing states  which,  in  man  at  least,  are  very 
different  in  subjective  quality.  For  this  reason  I 
am  inclined  to  urge  that  the  visceral  changes  mere- 
ly contribute  to  an  emotional  complex  more  or  less 
indefinite,  but  still  pertinent,  feelings  of  disturb- 
ance in  organs  of  which  we  are  not  usually  con- 
scious.    i\ 

This  view  that  the  differential  features  of  emo- 
tions are  not  to  be  traced  to  the  viscera  is  in  accord 


10 


with  the  experimental  results  of  Sherrington 
who  has  demonstrated  that  emotional  responses 
occur  in  dogs  in  which  practically  all  the  main  vis- 
cera and  the  great  bulk  of  skeletal  muscle  have 
been  removed  from  subjection  to  and  from  influ- 


INTERRELATIONS  OF  EMOTIONS    281 

ence  upon  the  brain,  by  severance  of  the  vagus 
nerves  and  the  spinal  cord.  In  these  animals  no 
alteration  whatever  was  noticed  in  the  occurrence, 
under  appropriate  circumstances,  of  characteristic 
expressions  of  voice  and  features,  indicating  anger, 
delight  or  fear.  The  argument  that  these  expres- 
sions may  have  been  previously  established  by  af- 
ferent impulses  from  excited  viscera  was  met  by 
noting  that  a  puppy  only  nine  weeks  old  also  con- 
tinued to  exhibit  the  signs  of  emotional  excitement 
after  the  brain  was  disconnected  from  all  the  body 
except  the  head  and  shoulders.  Evidence  from  uni- 
formity of  visceral  response  and  evidence  from  ex- 
clusion of  the  viscera  are  harmonious,  therefore,  in 
minimizing  visceral  factors  as  the  source  of  differ- 
ences in  emotional  states.* 

If  these  differences  are  due  to  other  than  vis- 
ceral changes,  why  is  it  not  always  possible  by  vol- 
untary innervations  to  produce  emotions  f  We  can 
laugh  and  cry  and  tremble.  But  forced  laughter 
does  not  bring  happiness,  nor  forced  sobbing  sor- 
row, and  the  trembling  from  cold  rouses  neither 
anger  nor  fear.  The  muscle  positions  and  tensions 
are  there,  but  the  experiencing  of  such  bodily 
changes  does  not  seem  even  approximately  to  rouse 

*  The  paucity  of  afferent  fibres  in  the  autonomic  system, 
and  the  probability  of  an  extremely  low  degree  of  sensitive- 
ness in  the  viscera  (for  evidence,  see  Cannon :  The  Mechan- 
ical Factors  of  Digestion,  London,  1911,  p.  202),  likewise  sup- 
port this  conclusion. 


282  BODILY   CHANGES 

an  emotion  in  us.  Voluntary  assumption  of  an  at- 
titude seems  to  leave  out  the  "feeling."  It  is  prob- 
able, however,  that  no  attitude  which  we  can  assume 
has  all  the  elements  in  it  which  appear  in  the  com- 
plete response  to  a  stirring  situation.  But  is  not  this 
because  the  natural  response  is  a  pattern  reaction, 
like  inborn  reflexes  of  low  order,  such  as  sneez- 
ing, in  which  impulses  flash  through  peculiarly  co- 
operating neurone  groups  of  the  central  system, 
suddenly,  unexpectedly,  and  in  a  manner  not  ex- 
actly reproducible  by  volition,  and  thus  they  throw 
the  skeletal  muscles  into  peculiar  attitudes  and,  if 
sufficiently  intense,  rush  out  in  diffuse  discharges 
that  cause  tremors  and  visceral  perturbations? 
The  typical  facial  and  bodily  expressions,  automat- 
ically assumed  in  different  emotions,  indicate  the 
discharge  of  peculiar  groupings  of  neurones  in  the 
several  affective  states.  That  these  responses  oc- 
cur instantly  and  spontaneously  when  the  appro- 
priate "situation,"  actual  or  vividly  imagined,  is 
present,  shows  that  they  are  ingrained  in  the  nerv- 
ous organization.  At  least  one  such  pattern,  that 
of  anger,  persists  after  removal  of  the  cerebral 
hemispheres — the  decorticated  dog,  by  growling 
and  biting  when  handled,  has  the  appearance  of 
being  enraged  ;^^  the  decerebrate  cat;  when  vig- 
orously stimulated,  retracts  its  lips  and  tongue, 
stares  with  dilated  pupils,  snarls  and  snaps  its 
jaws.^^     On  the  other  hand,   stroking  the  hair, 


IXTEEKELATIOXS  OF  EMOTIONS    283 

whistling  and  gently  calling  to  produce  a  pleased 
attitude,  or  yelling  to  produce  fright,  have  not  the 
slightest  effect  in  evoking  from  the  decorticated 
dog  signs  of  joy  and  affection  or  of  fear,  nor  does 
the  animal  manifest  any  sexual  feeling.  The  ab- 
sence of  bodily  indications  of  these  emotions  is 
quite  as  significant  as  the  presence  of  the  signs  of 
anger.  For  since  expressions  of  anger  can  persist 
without  the  cerebral  cortex,  there  is  little  reason 
why  the  complexes  of  other  emotional  expressions, 
if  their  "machinery"  exists  below  the  cortex,  should 
not  also  be  elicitable.  That  they  are  not  elicitable 
suggests  that  they  require  a  more  elaborately  or- 
ganized grouping  of  neurones  than  does  anger — 
possibly  what  the  cortex,  or  the  cortex  in  combi- 
nation with  basal  ganglia,  would  provide. 

The  contrast  between  the  brevity  of  the  "pseudo- 
affective  reactions"  in  the  decerebrate  cat,  though 
the  viscera  are  still  connected  with  the  central 
nervous  system,  and  the  normal  duration  of  emo- 
tional expression  in  the  dog  with  the  body  sepa- 
rated from  the  head  region,  has  been  used  by  Sher- 
rington to  weigh  the  importance  of  the  visceral  and 
other  factors.  And  the  evidence  which  I  have  given 
above,  as  well  as  that  which  he  has  offered,  favors 
the  view  that  the  viscera  are  relatively  unimpor- 
tant in  an  emotional  complex,  especially  in  con- 
tributing differential  features. 


284  BODILY    CHANGES 

REEERENCES 

1  James :  Principles  of  Psychology,  New  York,  1905,  i, 
p.  22. 

2  Prince :  The  Unconscious,  New  York,  1914,  p.  456. 

3  Hertz:  Constipation  and  Allied  Intestinal  Disorders, 
London,  1909,  p.  81. 

*v.  Trenck:  Merkwiirdige  Lebensgeschichte,  Berlin,  1787, 
p.  195. 

^  James,  Loc.  cit.,  p.  415. 

^  McDougall :  Introduction  to  Social  Psychology,  London, 
1908,  p.  72. 

'''  Darwin :  Loc.  cit.,  p.  76. 

s  Miiller :  Deutsches  Archiv  f iir  klinische  Medicin,  1907, 
Ixxxix,  p.  434. 

9  Burton :  The  Anatomy  of  Melancholy  (first  published  in 
1621),  London,  1886,  p.  443. 

1°  Sherrington :  Proceedings  of  the  Royal  Society,  1900, 
Ixvi,  p.  397. 

^^  Goltz :  Archiv  f iir  die  gesammte  Physiologie,  1892,  li, 
p.  577. 

^2  Woodworth  and  Sherrington :  Journal  of  Physiology, 
1904,  xxxi,  p.  234. 


CHAPTER   XV 

ALTERNATIVE  SATISFACTIONS  FOR  THE 
FIGHTING   EMOTIONS 

The  uniformity  of  visceral  responses  wlien  al- 
most any  feelings  grow  very  intense,  and  under 
such  conditions  the  identity  of  these  responses 
with  those  characteristically  aroused  in  the  bel- 
ligerent emotion  of  anger  or  rage  and  its  counter- 
part, fear,  offer  interesting  possibilities  of  trans- 
formation and  substitution.  This  is  especially 
true  in  the  activities  of  human  beings.  And  be- 
cause men  have  devised  such  terribly  ingenious  and 
destructive  modes  of  expressing  these  feelings  in 
war,  an  inquiry  into  the  basis  for  possible  substi- 
tution seems  not  out  of  place. 

Support  for  the  Militarist  Estimate  of  the  Strength  of 
THE  Fighting  Emotions  and  Instincts 

The  business  of  killing  and  of  avoiding  death 
has  been  one  of  the  primary  interests  of  living 
l)eings  throughout  their  long  history  on  the  earth. 
It  is  in  the  highest  degree  natural  that  feelings  of 

285 


286  BODILY   CHANGES 

hostility  often  burn  with  fierce  intensity,  and  then, 
with  astonishing  suddenness,  that  all  the  powers  of 
the  body  are  called  into  action — for  the  strength 
of  the  feelings  and  the  quickness  of  the  response 
measure  the  chances  of  survival  in  a  struggle 
where  the  issue  may  be  life  or  death.  These  are 
the  powerful  emotions  and  the  deeply  ingrained 
instinctive  reactions  which  invariably  precede  com- 
bat. They  are  the  emotions  and  instincts  that 
sometimes  seize  upon  individuals  in  groups  and 
spread  like  wildfire  into  larger  and  larger  aggre- 
gations of  men,  until  vast  populations  are  shout- 
ing and  clamoring  for  war.  To  whatever  extent 
military  plans  are  successful  in  devising  a  vast 
machine  for  attack  or  defense,  the  energies  that 
make  the  machine  go  are  found,  in  the  last  analysis, 
in  human  beings  who,  when  the  time  for  action 
comes,  are  animated  by  these  surging  elemental 
tendencies  which  assume  control  of  their  conduct 
and  send  them  madly  into  conflict. 

The  strength  of  the  fighting  instinct  in  man  has 
been  one  of  the  main  arguments  used  by  the  mili- 
tarists in  support  of  preparation  for  international 
strife.  They  point  to  the  historical  fact  that  even 
among  highly  civilized  peoples  scarcely  a  decade 
passes  without  a  kindling  of  the  martial  emo- 
tions, which  explode  in  actual  warfare.  Such  fight- 
ing, they  say,  is  inevitable — the  manifestation  of 
"biological  law" — and,  so  long  as  human  nature 


FIGHTING    EMOTIONS  287 

remains  unchanged,  decision  by  battle  must  be  re- 
sorted to.  They  urge,  furthermore,  that  in  war 
and  in  the  preparations  for  war  important  phys- 
ical qualities — sturdiness,  hardihood,  and  strength 
for  valorous  deeds — are  given  peculiarly  favorable 
opportunities  for  development,  and  that  if  these  op- 
portunities are  lacking,  lusty  youth  will  give  place 
to  weaklings  and  mollycoddles.  In  addition  the 
militarists  say  that  war  benefits  mankind  by  its 
moral  effects.  Without  war  nations  become  effete, 
their  ideals  become  tarnished,  the  people  sink  into 
self-indulgence,  their  wills  weaken  and  soften  in 
luxury.  War,  on  the  contrary,  disciplines  charac- 
ter, it  sobers  men,  it  teaches  them  to  be  brave  and 
patient,  it  renews  a  true  order  of  values,  and  its  de- 
mand for  the  supreme  sacrifice  of  life  brings  forth 
in  thousands  an  eager  response  that  is  the  crown- 
ing glory  of  the  human  spirit.  As  the  inevitable 
expression  of  a  deep-rooted  instinct,  therefore,  and 
as  a  unique  means  of  developing  desirable  physical 
and  moral  qualities,  war  is  claimed  by  the  mili- 
tarists to  be  a  natural  necessity.^ 

The  militarist  contention  that  the  fighting  in- 
stinct is  finally  fixed  in  human  nature  receives 
strong  confirmation  in  the  results  of  our  re- 
searches. Survival  has  been  decided  by  the  grim 
law  of  mortal  conflict,  and  the  mechanism  for  ren- 
dering the  body  more  competent  in  conflict  has  been 
revealed  in  earlier  chapters  as  extraordinarily  per- 


288  BODILY    CHANGES 

feet  and  complete.  Moreover,  the  physiological 
provisions  for  fierce  struggle  are  f onnd  not  only  in 
the  bodies  of  lower  animals,  that  must  hunt  and 
kill  in  order  to  live,  but  also  in  human  beings. 
Since  this  remarkable  mechanism  is  present,  and 
through  countless  generations  has  served  the  fun- 
damentally important  purpose  of  giving  momen- 
tous aid  in  the  struggle  for  existence,  the  mili- 
tarists might  properly  argue  that,  as  with  other 
physiological  processes,  bodily  harmony  would  be 
promoted  by  its  exercise.  Indeed,  they  might 
account  for  the  periodic  outburst  of  belligerent 
feelings  by  assuming  that  these  natural  aptitudes 
require  occasional  satisfaction.* 

Growing  Opposition  to  the  Fighting  Emotions  and  Instincts 
AS  Displayed  in  War 

In  spite  of  the  teachings  of  history  that  wars 
have  not  grown  fewer,  and  in  spite  of  the  militarist 
argument  that  war  is  a  means  of  purging  mankind 
of  its  sordid  vices,  and  renewing  instead  the  no- 
blest virtues,  the  conclusion  that  the  resort  to  arms 
is  unavoidable  and  desirable  is  nowadays  being 
strongly   contested.     The   militarists   show   only 

*  Mr.  Graham  Wallas  has  made  the  interesting  sugges- 
tion (The  Great  Society,  New  York,  1914,  p.  66)  that  nerv- 
ous strain  and  restlessness  due  to  "baulked  disposition"  may 
result  from  the  absence  of  circumstances  which  would  call 
the  emotional  responses  into  action.  And  he  cites  Aristotle's 
theory  that  pent  passions  may  be  released  by  represented 
tragedy  and  by  music. 


FIGHTING    EMOTIONS  289 

part  of  the  picture.  No  large  acquaintance  with 
the  character  of  warfare  is  necessary  to  prove  that 
when  elemental  anger,  hate  and  fear  prevail, 
civilized  conventions  are  abandoned  and  the  most 
savage  instincts  determine  conduct.  Homes  are 
looted  and  burned,  women  and  children  are 
abominably  treated,  and  many  innocents  are 
murdered  outright  or  starved  to  death.  No  bland 
argument  for  the  preservation  of  the  manly 
virtues  can  palliate  such  barbarities.  Even  when 
fighting  men  are  held  within  the  rules,  the  de- 
vices for  killing  and  injuring  are  now  made 
so  perfect  by  devilish  ingenuity  that  by  the 
pulling  of  a  trigger  one  man  can  in  a  few  seconds 
mow  down  scores  of  his  fellow-creatures  and  send 
them  writhing  to  agony  or  death.  War  has  become 
too  horrible;  it  is  conducted  on  too  stupendous  a 
scale  of  carnage  and  expenditure;  it  destroys  too 
many  of  the  treasured  achievements  of  the  race ;  it 
interferes  too  greatly  with  consecrated  efforts  to 
benefit  all  mankind  by  discovery  and  invention ;  it 
involves  too  much  suffering  among  peoples  not  di- 
rectly concerned  in  the  struggle ;  it  is  too  vastly  at 
variance  with  the  methods  of  fair  dealing  that  have 
been  established  between  man  and  man ;  the  human 
family  has  become  too  closely  knit  to  allow  some 
of  its  members  to  bring  upon  themselves  and  all 
the  rest  poverty  and  distress  and  a  long  heritage 
of  bitter  hatred  and  resolution  to  seek  revenge. 


290  BODILY    CHANGES 

All  these  reasons  for  hostility  to  war  imply  a 
thwarting  of  strong  desires  in  men — desires  for 
family  happiness,  devotion  to  beauty  and  to  schol- 
arship, passion  for  social  justice,  hopes  of  lessen- 
ing poverty  and  disease.  As  was  pointed  out  in 
the  previous  chapter,  the  feeling  of  hostility  has  no 
definite  object  to  awaken  it.  It  is  roused  when 
there  is  opposition  to  what  we  ardently  wish  to  get. 
And  because  war  brings  conditions  which  frustrate 
many  kinds  of  eagerly  sought  purposes,  war  has 
roused  in  men  a  hostility  against  itself.  There  is 
then  a  war  against  war,  a  willingness  to  fight 
against  monstrous  carnage  and  destruction,  that 
grows  in  intensity  with  every  war  that  is  waged. 

The  Desirability  of  Preserving  the  Martial  Virtues 

Although  there  is  increasing  opposition  to  the 
display  of  the  fighting  emotions  and  instincts  in 
war,  nevertheless  the  admirable  moral  and  phys- 
ical qualities,  claimed  by  the  militarists  to  be  the 
unique  products  of  war,  are  too  valuable  to  be  lost. 
As  McDougall  ^  has  indicated,  when  the  life  of 
ideas  becomes  richer,  and  the  means  we  take  to 
overcome  obstructions  to  our  efforts  more  refined 
and  complex,  the  instinct  to  fight  ceases  to  express 
itself  in  its  crude  natural  manner,  save  when  most 
intensely  excited,  and  becomes  rather  a  source  of 
increased  energy  of  action  towards  the  end  set  by 
any  other  instinct ;  the  energy  of  its  impulses  adds 


FIGHTING    EMOTIONS  291 

itself  to  and  reenforces  that  of  other  impulses  and 
so  helps  us  to  overcome  our  difficulties.  In  this 
lies  its  great  value  for  civilized  man.  A  man  de- 
void of  the  pugnacious  instinct  would  not  only  be 
incapable  of  anger,  but  would  lack  this  great  source 
of  reserve  energy  which  is  called  into  play  in  most 
of  us  by  any  difficulty  in  our  path. 

Thus  the  very  efficiency  of  a  war  against  war,  as 
well  as  struggle  against  other  evils  that  beset  civil- 
ized society,  rests  on  the  preservation  and  use  of 
aggressive  feeling  and  the  instinct  to  attack.  From 
this  point  of  view  the  insistence  by  the  militarists 
that  we  must  accept  human  nature  as  we  find  it, 
and  that  the  attempt  to  change  it  is  foolish,  seems 
a  more  justifiable  attitude  than  that  of  the  paci- 
fists who  belittle  the  fighting  qualities  and  urge 
that  changing  them  is  a  relatively  simple  process. 
We  should  not  wish  them  changed.  Even  if  in  the 
war  against  war  a  means  should  be  established  of 
securing  international  justice,  and  if  through  co- 
operative action  the  decrees  of  justice  were  en- 
forced, so  that  the  occasions  which  would  arouse 
belligerent  emotions  and  instincts  were  much  re- 
duced, there  would  still  remain  the  need  of  recog- 
nizing their  elemental  character  and  their  possible 
usefulness  to  society.  What  is  needed  is  not  a 
suppression  of  these  capacities  to  feel  and  act,  but 
their  diversion  into  other  channels  where  they  may 
have  satisfactory  expression. 


292  BODILY    CHANGES 

Moral  Substitutes  for  Warfare 

"We  must  make  new  energies  and  hardihoods 
continue  the  manliness  to  which  the  military  mind 
so  faithfully  clings.  Martial  virtues  must  be  the 
enduring  cement;  intrepidity,  contempt  of  soft- 
ness, surrender  of  private  interest,  obedience  to 
command,  must  still  remain  the  rock  upon  which 
states  are  built."  Thus  wrote  William  James  ^ 
in  proposing  a  "moral  equivalent  for  war."  This, 
he  suggested,  should  consist  of  such  required  serv- 
ice in  the  hard  and  difficult  occupations  as  would 
take  the  childishness  and  superciliousness  out  of 
our  youth  and  give  them  soberer  ideas  and  health- 
ier sympathies  with  their  fellow-men.  He  con- 
ceived that  by  proper  direction  of  its  education  a 
people  should  become  as  proud  of  the  attainment 
by  the  nation  of  superiority  in  any  ideal  respect 
as  it  would  be  if  the  nation  were  victorious  in  war. 
"The  martial  type  of  character,"  he  declared,  "can 
be  bred  without  war.  Strenuous  honor  and  disin- 
terestedness abound  elsewhere.  Priests  and  medi- 
cal men  are  in  a  fashion  educated  to  it,  and  we 
should  all  feel  some  degree  of  it  imperative  if  we 
were  conscious  of  our  work  as  an  obligatory  serv- 
ice to  the  state.  We  should  be  oivned,  as  soldiers 
are  by  the  army,  and  our  pride  would  rise  ac- 
cordingly. We  could  be  poor,  then,  without 
humiliation,  as  army  officers  now  are.  The  only 
thing  needed  henceforth  is  to  inflame  the  civic 


FIGHTING    EMOTIONS  293 

temper  as  past  history  has  inflamed  the  military 
temper." 

Similar  ideas  have  been  expressed  by  others."* 
It  has  been  pointed  out  that  the  great  war  of  man- 
kind is  that  against  pain,  disease,  poverty  and  sin ; 
that  the  real  heroes  are  not  those  who  squander  hu- 
man strength  and  courage  in  fighting  one  another, 
but  those  who  fight  for  man  against  these  his  eter- 
nal foes.  War  of  man  against  man,  in  this  view, 
becomes  dissension  in  the  ranks,  permitting  the 
common  enemies  to  strike  their  most  telling  blows. 

These  moral  considerations,  however,  are  apart 
from  the  main  intent  of  our  discussion.  Our  ear- 
lier inquiry  confirmed  the  belief  that  the  fighting 
emotions  are  firmly  rooted  in  our  natures,  and 
showed  that  these  emotions  are  intimately  asso- 
ciated with  provisions  for  physical  exertion.  It 
is  particularly  in  this  aspect  of  the  discussion  of 
substitutes  for  war  that  these  studies  have  sig- 
nificance. 

Physical  Substitutes  for  Warfare 

The  idealization  of  the  state  and  the  devotion 
of  service  to  social  welfare,  which  have  been  sug- 
gested as  moral  substitutes  for  military  loyalty, 
leave  unanswered  the  claims  of  the  militarists 
that  in  war  and  in  preparations  for  war  oppor- 
tunities are  offered  which  are  peculiarly  favorable 
to  the  development  of  important  physical  qualities 


294  BODILY    CHANGES 

— bodily  vigor,  sturdiness,   and  ability  to  with- 
stand all  manner  of  hardships. 

In  the  evidence  previously  presented,  it  seems 
to  me  there  was  a  suggestion  that  offers  a  perti- 
nent alternative  to  these  claims.  When  the  body 
goes  onto  what  we  have  called  a  war  footing,  the 
physiological  changes  that  suddenly  occur  are 
all  adapted  to  the  putting  forth  of  supreme  mus- 
cular and  nervous  efforts.  That  was  what  primi- 
tive battle  consisted  of,  through  countless  myriads 
of  generations — a  fierce  physical  contest  of  beast 
with  beast,  and  of  man  with  man.  Such  contests, 
attended  as  they  were  by  the  thrill  of  unpredict- 
able incidents,  and  satisfying  completely  the  lust 
of  combat,  are  to  be  contrasted  with  the  dull  grind 
in  preparation  for  modern  war,  the  monotonous 
regularity  of  subservience,  the  substitution  every- 
where of  mechanism  for  muscle,  and  often  the 
attack  on  an  enemy  who  lies  wholly  unseen,*    As 

*Lord  Wolseley,  while  commander-in-chief  of  the  English 
forces,  in  1897,  secured  sanction  for  not  displaying  the  regi- 
mental colors  in  battle.  "It  would  be  madness  and  a  crime," 
he  declared,  "to  order  any  soldier  to  carry  colors  into  action 
in  the  future.  You  might  quite  as  well  order  him  to  be  as- 
sassinated. We  have  had  most  reluctantly  to  abandon  a 
practice  to  which  we  attached  great  importance,  and  which, 
under  past  and  gone  conditions  of  fighting,  was  invaluable  in 
keeping  alive  the  regimental  spirit  upon  which  our  British 
troops  depended  so  much."  All  was  has  been  transformed  by 
the  invention  of  the  far-reaching  and  fate-dealing  rifle  and 
automatic  gun,  with  which  an  enemy  kills,  whose  face  is  not 
even  seen.     War  is  almost  reduced  to  a  mechanical  inter- 


FIGHTING   EMOTIONS  295 

Wallas  with  nice  irony  has  remarked,  "The  gods 
in  Valhalla  would  hardly  choose  the  organization 
of  modem  lines  of  military  communication,  as 
they  chose  the  play  of  sword  and  spear,  to  be  the 
most  exquisite  employment  of  eternity." 

While  it  is  true  that  physical  strength  can  be 
developed  by  any  form  of  hard  labor,  as,  for 
example,  by  sawing  wood  or  digging  ditches,  such 
labor  does  not  stimulate  quickness,  alertness,  and 
resourcefulness  in  bodily  action.  Nor  does  it  give 
any  occasion  for  use  of  the  emotional  mechanism 
for  reenforcement.  If  this  mechanism,  like  other 
physiological  arrangements,  is  present  in  the 
body  for  use — and  iDrevious  discussion  leaves  little 


change  of  volleys  and  salvoes,  and  to  the  intermittent  fire  of 
rifles  and  machine  guns,  with  short  rushes  at  the  last,  in 
which  there  is  no  place  for  the  dignity  and  grace  of  the 
antique  battle  of  the  standard.  (See  London  Times,  July  31, 
1897,  p.  12.) 

T.  F.  Millard,  the  well-known  correspondent  of  the  Russo- 
Japanese  War,  wrote  as  follows  of  the  characteristics  of 
present  day  conflicts :  "A  large  part  of  modern  war  is  on 
too  great  a  scale  to  give  much  opportunity  for  individual 
initiative.  Soldiers  can  rarely  tell  what  is  going  on  in  their 
immediate  vicinity.  They  cannot  always  see  the  enemy  they 
are  firing  at,  and  where  they  can  see  the  object  of  their  fire 
such  an  important  matter  as  range  and  even  direction  can- 
not be  left  to  them.  .  .  .  Troops  are  clothed  so  much  alike 
nowadays  that  it  is  very  difficult  to  distinguish  friend  from 
foe  at  five  hundred  yards,  and  large  bodies  of  troops  rarely 
get  that  close  to  each  other  in  modern  war  while  there  is 
light  enough  to  see  clearly.  .  .  .  Battery  officers  simply  see 
that  their  guns  are  handled  according  to  instructions.    They 


296  BODILY   CHANGES 

doubt  of  that — then  as  a  means  of  exercising  it 
and,  in  addition,  satisfying  the  strong  instinct  for 
competitive  testing  of  strength  and  physical  skill, 
some  activity  more  enlivening  than  monotonous 
gymnastics  and  ordered  marching  is  required. 

In  many  respects  strenuous  athletic  rivalries 
present,  better  than  modern  military  service,  the 
conditions  for  which  the  militarists  argue,  the 
conditions  for  which  the  body  spontaneously  pre- 
pares when  the  passion  for  fighting  prevails.  As 
explained  in  an  earlier  chapter,  in  competitive 
sports  the  elemental  factors  are  retained — man 
is  again  pitted  against  man,  and  all  the  resources 
of  the  body  are  summoned  in  the  eager  struggle 


regulate  the  time,  speed,  objective  and  range  as  ordered.  .  .  . 
The  effects  of  the  fire  are  observed  by  officers  appointed  to 
that  duty,  stationed  at  various  parts  of  the  field,  often  miles 
and  miles  apart,  and  who  are  in  constant  communication 
with  the  chief  of  artillery  by  telephone."  (See  Scribner's 
Magazine,  1905,  xxxvii,  pp.  64,  66.) 

The  testimony  of  a  captain  of  a  German  battery  engaged 
against  the  French  and  English  in  1914,  supports  the  forego- 
ing claims.  He  is  reported  as  saying :  "We  shoot  over  those 
tree  tops  yonder  in  accordance  with  directions  for  range  and 
distance  which  come  from  somewhere  else  over  a  field  tele- 
phone, but  we  never  see  the  men  at  whom  we  are  firing. 
They  fire  back  without  seeing  us,  and  sometimes  their  shells 
fall  short  or  go  beyond  us,  and  sometimes  they  fall  among  us 
and  kill  and  wound  a  few  of  us.  Thus  it  goes  on  day  after 
day.  I  have  not  with  my  own  eyes  seen  a  Frenchman  or  an 
Englishman  unless  he  was  a  prisoner.  It  is  not  so  much 
pleasure — fighting  like  this."  (See  Philadelphia  Saturday 
Evening  Post,  December  26,  1914,  p.  27.) 


FIGHTING    EMOTIONS  297 

for  victory.  And  because,  under  such  circum- 
stances, the  same  physiological  alterations  occur 
that  occur  in  anticipation  of  mortal  combat,  the 
belligerent  emotions  and  instincts,  so  far  as  their 
bodily  manifestations  are  concerned,  are  thereby 
given  complete  satisfaction. 

The  Significance  of  International  Athletic  Competitions 

For  reasons  given  above,  I  venture  to  lay  em- 
phasis on  a  suggestion,  which  has  been  made 
before  by  others,  that  the  promotion  of  great  inter- 
national athletic  contests,  such  as  the  Olympic 
games,  would  do  for  our  young  men  much  that  is 
now  claimed  as  peculiar  to  the  values  of  military 
discipline.  The  substitution  of  athletic  rivalries 
for  battle  is  not  unknown.  In  the  Philippine  Isl- 
ands, according  to  Worcester,^  there  were  no 
athletics  before  the  American  occupation.  The 
natives  soon  learned  games  from  the  soldiers. 
And  when  the  sports  reached  such  development 
that  comi^etition  between  towns  and  provinces  was 
possible,  they  began  to  arouse  the  liveliest  enthu- 
siasm among  the  people.  The  physical  develop- 
ment of  the  participants  has  been  greatly  stimu- 
lated, the  spirit  of  fair  play  and  sportsmanship, 
formerly  lacking,  has  sprung  into  existence  in 
every  section  of  the  Islands,  and  the  annual  meets 
between  athletic  teams  from  various  provinces  are 
recognized  as  promoting  a  general  and  friendly 


298  BODILY    CHANGES 

understanding  among  the  different  Filipino  tribes. 
The  fierce  Igarots  of  Bontoc,  once  constantly  at 
war  with  neighboring  tribes,  now  show  their  prow- 
ess not  in  head-himting,  but  in  baseball,  wrestling, 
and  the  tug-of-war.* 

Is  it  unreasonable  to  expect  that  what  has  hap- 
pened in  the  Philippine  Islands  might,  by  proper 
education  and  suggestion,  happen  elsewhere  in  the 
world!  Certainly  the  interest  in  athletic  contests 
is  no  slight  and  transient  interest.  At  the  time 
of  a  great  war  we  know  that  news  of  the  games 
is  fully  as  much  demanded  as  news  of  the  war. 
Already  in  the  United  States,  without  special 
stimulation,  the  number  of  young  men  engaged  in 
athletic  training  is  estimated  as  equal  to  the  num- 
ber in  the  standing  army.  And  in  England,  belief 
in  the  efficacy  of  athletics  as  a  means  of  promoting 
hardihood  and  readiness  to  face  stern  hazards  has 
found  expression  in  the  phrase  that  England's 
battles  have  been  won  on  the  football  fields  of 
Eugby  and  of  Eton.  With  the  further  promotion 
of  international  contests  the  influence  of  competi- 
tive sports  is  likely  to  increase  rather  than  lessen. 
Within  national  boundaries  emulation  is  sure  to 
stimulate  extensively  such  games  as  will  bring 
forth  the  best  representative  athletes  that  the  coun- 

*  It  is  reported  that  when  these  warriors  first  appeared  at 
the  games,  each  brought  his  spear,  which  he  drove  into  the 
ground  beside  him,  ready  for  use.  As  the  nature  of  the  new 
rivalries  became  known,  the  spears  were  left  behind. 


FIGHTING   EMOTIONS  299 

try  can  produce.  In  one  of  the  high-spirited  Eu- 
ropean nations,  which  made  a  poor  showing  at  the 
last  Olympic  meet,  thousands  of  young  men  began 
training  for  the  next  meet,  under  a  director  im- 
ported from  the  nation  that  had  made  the  highest 
records. 

Training  for  athletic  contests  is  quite  as  likely 
to  enure  young  men  to  physical  hardship  and 
fatigue,  is  quite  as  conducive  to  the  development 
of  bodily  vigor,  the  attainment  of  alertness  and 
skill  and  the  practice  of  self-restraint,  as  is  army 
life  with  its  traditional  associations  and  easy  li- 
cense. It  may  be  urged,  however,  that  an  essential 
element  is  lacking  in  all  this  discussion — the  so- 
bering possibility  that  in  war  the  supreme  sur- 
render of  life  itself  may  be  required.  Death  for 
one's  country  is  indeed  glorious.  But  the  argu- 
ment that  being  killed  is  desirable  has  little  to 
commend  it.  When  the  strongest  and  sturdiest  are 
constantly  chosen  to  be  fed  to  the  engines  of  anni- 
hilation, the  race  is  more  likely  to  lose  greater 
values  than  it  gains  from  the  spectacle  of  self- 
sacrifice,  however  perfect  that  may  be.  Are  there 
not  advantages  in  the  conditions  of  great  athletic 
rivalries  that  may  compensate  for  war's  most  aus- 
tere demand?  The  race  of  hardy  men,  to  secure 
which  the  militarists  urge  war,  is  much  more  likely 
to  result  from  the  honoring  and  preserving  of 
vigorous  men  in  their  vigor  than  it  is  from  the 


300  BODILY    CHANGES 

systematic  selection  of  such  men  to  be  destroyed 
in  their  youth. 

There  are  other  aspects  of  international  games 
which  strongly  commend  them  as  an  alternative 
to  the  pursuit  of  military  discipline.  The  high 
standards  of  honor  and  fairness  in  sport ;  its  un- 
failing revelation  of  excellence  without  distinc- 
tions of  class,  wealth,  race  or  color ;  the  ease  with 
which  it  becomes  an  expression  of  the  natural 
feelings  of  patriotism;  the  respect  which  victory 
and  pluckily  borne  defeat  inspire  in  competitors 
and  spectators  alike;  the  extension  of  acquaint- 
ance and  understanding  which  follows  from 
friendly  and  magnanimous  rivalry  among  strong 
men  who  come  together  f-rom  the  ends  of  the  earth 
— each  of  these  admirable  features  of  athletic  con- 
tests between  nations  might  be  enlarged  upon. 
But,  as  intimated  before,  these  moral  considera- 
tions must  be  left  without  further  mention,  as 
being  irrelevant  to  the  physiological  processes 
with  which  we  are  dealing. 

We  are  concerned  with  the  question  of  exercis- 
ing the  fighting  instinct  and  thus  assuring  the 
physical  welfare  of  the  race.  The  race  must  de- 
generate, the  militarists  say,  if  this  instinct  is 
not  allowed  to  express  itself  in  war.  This  declar- 
ation we  are  in  a  position  to  deny,  for  the  evi- 
dence is  perfectly  clean-cut  that  the  aggressive 
instincts,  which  through  aeons  of  racial  experi- 


FIGHTING   EMOTIONS  301 

ence  liave  naturally  and  spontaneously  developed 
vigor  and  resourcefulness  in  the  body,  are  invited 
by  elemental  emotions,  and  that  through  these 
emotions  energies  are  released  which  are  highly 
useful  to  great  physical  effort.  No  stupid  routine 
of  drill,  or  any  other  deadening  procedure,  will 
call  these  energizing  mechanisms  into  activity. 
War  and  the  preparations  for  war  nowadays  have 
become  too  machine-like  to  serve  as  the  best  means 
of  preserving  and  disciplining  these  forces.  The 
exhilarating  swing  and  tug  and  quick  thrust  of  the 
big  limb  muscles  have  largely  vanished.  Pressing 
an  electric  contact  or  bending  the  trigger  finger 
is  a  movement  altogether  too  trifling.  If,  then, 
natural  feelings  must  be  expressed,  if  the  fight- 
ing functions  of  the  body  must  be  exercised, 
how  much  better  that  these  satisfactions  be  found 
in  natural  rather  than  in  artificial  actions,  how 
much  more  reasonable  that  men  should  struggle 
for  victory  in  the  ancient  ways,  one  against  an- 
other, body  and  spirit,  as  in  the  great  games. 

REFERENCES 

^  See  Angell:  The  Great  Illusion,  New  York  and  London, 

1913,  pp.  159-1G4. 

2  McDougall :  Introduction  to  Social  Psychology,  London, 
1908,  p.  01. 

^  James:  Memories  and  Studies,  New  York,  1911,  p.  287. 

*  See  Perry : :  The  Moral  Economy,  New  York,  1909,  p.  32 ; 
and  Drake:  Problems  of  Conduct,  Boston,  1914,  p.  317. 

°  Worcester :  The  Philippines,  Past  and  Present,  New  York, 

1914,  ii,  pp.  515,  578. 


A  LIST  OF  PUBLISHED  KESEAECHES  EEOM  THE 
PHYSIOLOGICAL  LABORATORY  IN  HARVARD 
'     UNIVERSITY,   ON   WHICH   THE   PRES- 
ENT ACCOUNT  IS  BASED. 

1.  The  Influence  of  Emotional  States  on  the  Functions  of 
the  Alimentary  Canal.  By  W.  B.  Cannon.  American  Jour- 
nal of  the  Medical  Sciences,  1909,  cxxxvii,  pp.  480-487. 

2.  Emotional  Stimulation  of  Adrenal  Secretion.  By  W.  B. 
Cannon  and  D.  de  la  Paz.  American  Journal  of  Physiology, 
1911,  xxviii,  pp.  64-YO. 

3.  The  Effects  of  Asphyxia,  Hyperpnoea,  and  Sensory 
Stimulation  on  Adrenal  Secretion.  By  W.  B.  Cannon  and 
R.  G.  Hoskins.    Ihid.,  1911,  xxix,  pp.  274-279. 

4.  Emotional  Glycosuria.  By  W.  B.  Cannon,  A.  T.  Shohl 
and  W.  S.  Wright.    Ihid.,  1911,  xxix,  pp.  280-287. 

5.  A  Consideration  of  Some  Biological  Tests  for  Epi- 
nephrin.  By  R.  G.  Hoskins.  Journal  of  Pharmacology  and 
Experimental  Therapeutics,  1911,  iii,  pp.  93-99. 

6.  The  Sthenic  Effect  of  Epinephrin  upon  Intestine.  By 
R.  G.  Hoskins.  American  Journal  of  Physiology,  1912,  xxix, 
pp.  363-366. 

7.  An  Explanation  of  Hunger.  By  W.  B.  Cannon  and  A. 
L.  Washburn.    Ihid.,  1912,  xxix,  pp.  441-454. 

8.  A  New  Colorimetric  Method  for  the  Determination  of 
Epinephrin.  By  O.  Folin,  W.  B.  Cannon  and  W.  Denis. 
Journal  of  Biological  Chemistry,  1913,  xiii,  pp.  477-483. 

9.  The  Depressor  Effect  of  Adrenalin  on  Arterial  Pressure. 
By  W.  B.  Cannon  and  Henry  Lyman.  American  Journal  of 
Physiology,  1913,  xxxi,  pp.  376-398. 

302 


PUBLISHED    RESEARCHES  303 

10.  The  Effect  of  Adrenal  Secretion  on  Muscular  Fatigue. 
By  W.  B.  Cannon  and  L.  B.  Nice.  Ihid.,  1913,  xxxii,  pp. 
44-60. 

11.  Fatigue  as  Affected  by  Changes  of  Arterial  Pressure. 
By  C.  M.  Gruber.    Ibid.,  1913,  xxxii,  pp.  222-229. 

12.  The  Threshold  Stimulus  as  Affected  by  Fatigue  and 
Subsequent  Best.  By  C.  M.  Gruber.  Ibid.,  1913,  xxxii,  pp. 
438-449. 

13.  The  Fatigue  Threshold  as  Affected  by  Adrenalin  and 
by  Increased  Arterial  Pressure.  By  C.  M.  Gruber.  Ibid., 
1914,  xxxiii,  pp.  335-355. 

14.  The  Emergency  Function  of  the  Adrenal  Medulla  in 
Pain  and  the  Major  Emotions.  By  W.  B.  Cannon.  Ibid., 
1914,  xxxiii,  pp.  356-372. 

15.  The  Relation  of  Adrenalin  to  Curare  and  Fatigue  in 
Normal  and  Denervated  Muscles.  By  C.  M.  Gruber.  Ibid., 
1914,  xxxiv,  pp.  89-96. 

16.  The  Graphic  Method  of  Recording  Coagulation.  By 
\V.  B.  Cannon  and  W.  L.  Mendenhall.  Ibid.,  1914,  xxxiv, 
pp.  225-231. 

17.  The  Hastening  or  Retarding  of  Coagulation  by 
Adrenalin  Injections.  By  W.  B.  Cannon  and  Horace  Gray. 
Ibid.,  1914,  xxxiv,  pp.  232-242. 

18.  The  Hastening  of  Coagulation  by  Stimulating  the 
Splanchnic  Nerves.  By  W.  B.  Cannon  and  W.  L.  Menden- 
hall.   Ibid.,  1914,  xxxiv,  pp.  243-250. 

19.  The  Hastening  of  Coagulation  in  Pain  and  Emotional 
Excitement.  By  W.  B.  Cannon  and  W.  L.  Mendenhall. 
Ibid.,  1914,  xxxiv,  pp.  251-261. 

20.  The  Interrelations  of  Emotions  as  Suggested  by  Recent 
Physiological  Researches.  By  W.  B.  Cannon.  American 
Journal  of  Psychology,  1914,  xxv,  pp.  256-282. 


INDEX 


I 


Adrenal  extract :  effect  of, 
on  muscular  contraction, 
82. 

Adrenal  glands :  nerve  sup- 
ply of,  37;  stimulated  in 
emotion,  52-59,  62-G3 ; 
stimulated  in  pain,  59-62, 
63;  in  relation  to  blood 
sugar,  77 ;  removal  of, 
causes  muscular  weakness, 
81 ;  secretion  of,  improves 
contraction  of  fatigued 
muscle,  92;  variations  in 
adrenin  content  of,  171 ; 
latent  period  of,  vphen 
splanchnics  stimulated, 
188 ;  amount  of  secretion 
from,  when  splanchnics 
stimulated,  198 ;  fatigue  of, 
199;  stimulated  by  as- 
phyxia, 206-208. 

Adrenin :  secreted  by  adre- 
nal glands,  36 ;  action  of, 
identical  with  sympathetic 
impulses,  37,  64;  secretion 
of,  by  splanchnic  stimula- 
tion, 41-43;  secreted  in 
emotional  excitement,  44, 
52-59;  method  of  testing 
for,  in  blood,  47-50 ;  se- 
creted in  emotion,  52-59, 
G2-C3;  disappearance  of, 
from  blood,  58 ;  secreted  in 
pain,  59-62,  63;  effects  of, 
when  injected  into  body, 
64-65;  effect  of,  on  dis- 
tribution   of    blood   in    tlie 


body,  107 ;  quickly  restores 
fatigued  muscle  to  normal 
irritability,  119-123 ;  specific 
in  its  restorative  action, 
124-128 ;  as  an  antidote  to 
muscular  metabolites,  129; 
restores  fatigued  denerv- 
ated  muscle  to  normal  irri- 
tability, 130;  point  of  ac- 
tion of,  in  muscle,  128-133; 
antagonistic  to  curare,  132 ; 
induces  rapid  coagulation 
of  blood,  136,  147  ff.;  not 
the  direct  cause  of  rapid 
coagulation,  156-158;  fails 
to  shorten  coagulation  time 
in  absence  of  intestines 
and  liver,  157-158;  vari- 
able amount  of,  in  adrenal 
glands,  171 ;  emergency 
functions  of,  185  tf . ;  util- 
ity of,  in  bettering  the  con- 
traction of  fatigued  mus- 
cle, 194-195;  not  a  check 
to  use  of  sugar  in  the  body, 
197,  199;  amount  of,  se- 
creted when  splanchnics 
stimulated,  198;  a  condition 
for  increase  of  blood  sugar, 
199;  stimulates  the  heart, 
191,  201 ;  dilates  the  bron- 
chioles, 204;  secretion  of, 
increased  in  asphyxia,  206- 
208. 
Amyl  nitrite:  effect  of,  on 
contraction  of  fatigued 
muscle,  126. 


305 


306 


INDEX 


Anger:  associated  with  ac- 
tion, 188 ;  energizing  influ- 
ence of,  216. 

Antagonisms :  autonomic,  34 ; 
in  relation  to  emotions,  38 ; 
between  cranial  and  sym- 
pathetic divisions,  268-270; 
between  sacral  and  sympa- 
thetic divisions,  270-272. 

Appetite :  compared  with 
hunger,,  233,  235;  opera- 
tion of,  after  section  of 
vagus  and  splanchnic 
nerves,  240. 

Arterial  blood  pressure:  in- 
creased in  excitement,  95; 
artificial  methods  of  in- 
creasing, 97;  influence  of 
different  heights  of,  on 
fatigue,  97-102 ;  influence 
of  increase  of,  on  fa- 
tigue, 97-102 ;  influence 
of  decrease  of,  on  fa- 
tigue, 102  -  104;  the 
"critical  region"  in  de- 
creasing, 104;  explanation 
of  effects  on  fatigued  mus- 
cle, of  varying,  104-106; 
value  of  increased,  in  pain  ' 
and  emotion,  106. 

Arteries :  innervation  of,  26. 

Asphyxia :  increases  adrenal 
secretion,  206-208 ;  in- 
creases sugar  in  blood,  209. 

Athletes :  glycosuria  of,  after 
games,  75. 

Autonomic  nervous  system: 
three  divisions  of,  25;  ar- 
rangement of  sympathetic 
division  of,  26-29 ;  arrange- 
ment of  cranial  and  sacral 
divisions  of,  29-30;  general 
functions  of  cranial  di- 
vision of,  30-32;  general 
functions  of  sacral  division 
of,  32-34;  antagonism  be- 
tween sympathetic  and  ' 
cranial-sacral  divisions  of. 


34-36 ;  identity  of  action  of 
sympathetic  division  of, 
and  adrenal  secretion,  36- 
38 ;  antagonisms  between 
emotions  expressed  in,  268- 
272. 

Behavior :  biological  explana- 
tion of,  2. 

Bile:  flow  of,  inhibited  by 
excitement,  13. 

Bladder:  innervation  of,  27, 
32;  effects  of  emotions  on, 
33. 

Blood :  method  of  obtaining, 
for  test  for  adrenin,  45-46; 
method  of  testing,  for 
adrenin,  47-50;  sugar  in, 
66,  73-74;  distribution  of, 
as  affected  by  adrenin,  pain 
and  excitement,  107-108, 
200;  functions  of,  135; 
rapid  coagulation  of,  by 
adrenin,  136  ff. ;  drawing 
of,  for  testing  coagulation 
time,  140-142 ;  treatment 
of,  in  testing  coagulation 
time,  142-145;  faster  co- 
agulation of,  after  subcu- 
taneous injections  of  adre- 
nin, 147-150,  and  after  in- 
travenous injections,  150- 
156;  oscillations  in  the 
rate  of  coagulation  of, 
155 ;  rapid  coagulation 
of,  not  due  directly  to  ad- 
renin, 156-158;  rapid  co- 
agulation of,  not  caused  by 
adrenin  in  absence  of  liver 
and  intestines,  157-158, 
and  not  caused  by  increase 
of  blood  sugar,  159,  170; 
coagulation  of,  hastejied 
by  splanchnic  stimulation, 
162-167,  but  not  in  absence 
of  adrenal  glands,  167-171 ; 
possible  delay  of  coagula- 
tion  of,   after   stimulation 


INDEX 


307 


of  hepatic  nerves,  170;  co- 
agulation of,  hastened  by 
"painful"  stimulation,  172- 
177 ;  coagulation  of,  hasten- 
ed in  light  anesthesia,  174- 
177 ;  rapid  coagulation  of, 
after  excitement,  stopped 
by  severing  splanchnic 
nerves,  180-182;  utility  of 
increased  sugar  in,  188-193  ; 
distribution  of,  in  pain 
and  excitement,  favorable 
to  muscular  effort,  201 ; 
sugar  in,  increased  by  as- 
phyxia, 209 ;  utility  of 
rapid  coagulation   of,  211. 

Bronchioles  :  dilated  by  adre- 
nin,  204. 

Bulimia :  explanation  of, 
262. 

Coagulation,   see  Blood. 

Coagulometer :  graphic,  138- 
147. 

Combat:  relation  of  emotion 
and  endurance  in,  225- 
226;  nature  of  ancient, 
294. 

Constipation:  as  result  of 
worry  and  anxiety,  271. 

Cortex,  cerebral :  insensitive- 
ness  of,  242. 

Cranial  autonomic  division; 
functions  of,  to  conserve 
bodily  resources,  30-32, 
268;  activities  of,  suppress- 
ed by  activities  of  sym- 
pathetic division,  268- 
272. 

Curare :  action  of,  antago- 
nized by  adrenin,  132. 

Dances :  relation  of  excite- 
ment and  endurance  in, 
222-224. 

Danger :  stimulating  effect 
of,  230. 

Dervishes:  exhibitions  of  en- 
durance by,  224. 


Digestion :  interruption  of, 
by  strong  emotion,  9-12, 
13-18,  268-269. 

Emotions :  surface  signs  of, 
3 ;  favorable  to  digestive  se- 
cretions, 4-8;  unfavorable 
to  digestive  secretions,  9- 
13 ;  persistence  of  effects 
of,  on  digestive  secretions, 
12;  effects  of,  on  gastric 
and  intestinal  contractions, 
13-18 ;  in  relation  to  sym- 
pathetic division,  36  ;•  in  re- 
lation to  adrenal  secretion, 
44,  52-59,  62-63;  increase 
of  blood  sugar  in,  66,  73 ; 
glycosuria  in,  70-76;  influ- 
ence of,  on  distribution  of 
blood  in  body,  108 ;  faster 
coagulation  of  blood  in, 
177-182,  but  stopped  by 
cutting  splanchnics,  180- 
182 ;  value  of  forced  res- 
piration in,  203 ;  value  of 
bronchiolar  dilation  in, 
204;  relation  to  action,  215  ; 
displayed  in  a  "pattern" 
response,  218,  282 ;  in  re- 
lation to  exhibitions  of 
power  and  endurance,  215, 
229 ;  antagonisms  between 
cranial  and  sympathetic, 
268-270,  and  between  sacral 
and  sympathetic,  270- 
272 ;  similarity  of  visceral 
changes  in  strong,  275-279  ; 
dependence  of,  on  cerebral 
cortex,  282-283. 

Endurance :  feats  of,  related 
to  great  emotion,  217-218; 
in  the  excitements  of 
mania  and  dancing,  222- 
224;  stimulated  by  music, 
228. 

Esophagus :  contractions  of, 
associated  with  hunger  sen- 
sation, 259-260. 


308 


INDEX 


Fatigue :  of  muscle,  84 ;  mus- 
cular, lessened  by  splanch- 
nic stimulation,  89-93;  as 
affected  by  increase  of  ar- 
terial pressure,  97-103 ; 
irritability  of  muscle  in, 
increased  by  splanchnic 
stimulation,  101;  explana- 
tion of  effects  of  varied  ar- 
terial pressure  on,  104-106 ; 
lessens  neuro-muscular  ir- 
ritability, 114-117,  120 ;  ef- 
fect of,  on  curarized  mus- 
cle, 132;  utility  of  adrenin 
in  lessening  effects  of,  194, 
195;  of  adrenal  glands, 
199 ;  cessation  of  hunger 
contractions  in,  262. 

Fear:  anticipatory  character 
of,  186-187 ;  associated  with 
action,  188 ;  explanation 
of  paralyzing  effect  of, 
189 ;  energizing  influence 
of,  216;  relation  to  rage, 
276;  bodily  changes  in, 
like  those  in  rage,  276-277 ; 
importance  of,  as  a  fight- 
ing emotion,  286. 

'Tesselungsdiabetes,"  69. 

Fever :  absence  of  hunger  in, 
242,  263. 

Fighting  emotions :  bodily 
changes  in,  like  those  in 
competitive  sports,  219-221, 
296;  anger  and  fear  as, 
285;  importance  of,  286; 
satisfactions  for,  in  com- 
petitive sports,  301. 

Food:  effect  of  sight  and 
smell  of,  on  gastric  secre- 
tion, 6. 

Football:  glycosuria  in  play- 
ers of,  75;  relation  of  ex- 
citement and  power  in, 
219-221. 

Frenzy:  endurance  in,  223, 
224. 


Ganglia :    autonomic,   23. 

Gastric  glands :  turgescence 
of,  not  the  cause  of  hun- 
ger sensation,  249-250. 

Gastric  juice :  psychic  secre- 
tion of,  5-8,  11 ;  importance 
of,  for  intestinal  digestion, 
7 ;  flow  of,  inhibited  by  ex- 
citement, 9-12,  and  by 
pain,  19. 

Generative  organs:  innerva- 
tion of,  32,  33;  effects  of 
strong  emotions  on  activ- 
ities in,  271. 

Glycosuria:  in  pain,  69-70; 
in  emotion,  70-76;  after 
football,  75,  221;  after  ex- 
aminations, 76 ;  depend- 
ence of,  on  adrenal  glands, 
77. 

Heart :  innervation  of,  26,  31 ; 
use  of  sugar  by,  191; 
stimulated  by  adrenin, 
191,  201. 

Hunger:  compared  with  appe- 
tite, 233,  235;  description 
of,  234-236;  theories  of, 
237 ;  as  a  general  sensation, 
237;  disappearance  of,  as 
time  passes,  238-239 ;  when 
stomach  full,  239;  may  be 
absent  in  bodily  need,  242- 
243 ;  temporarily  abolished 
by  indigestible  materials, 
243;  quick  onset  and  peri- 
odicity of,  244-245;  refer- 
ence of,  to  stomach  region, 
245-247;  not  due  to  empti- 
ness of  stomach,  248;  not 
due  to  hydrochloric  acid  in 
empty  stomach,  248;  not 
due  to  turgid  gastric 
glands,  249-250;  as  the  re- 
sult of  contractions,  251- 
253;  inhibited  by  swallow- 
ing, 254 ;  method  of  record- 
ing gastric  contractions  in. 


INDEX 


309 


255-256 ;  associated  with 
gastric  contractions,  256- 
259,  and  with  esophageal 
contractions,  259  -  260; 
function  of,  263-264,  272- 
275. 
Hydrochloric  acid :  not  the 
cause  of  hunger  sensation, 
248. 

Intestine :  contractions  of, 
inhibited  by  excitement, 
16;  innervation  of,  27,  31; 
use  of,  as  test  for  adrenin 
in  blood,  47-50;  contracts 
when  empty,  251-253  ;  con- 
tractions of,  may  originate 
hunger  sensations,  263. 

Instincts:  relation  of, to  emo- 
tions, 187,  188. 

Irritability:  increased  in  fa- 
tigued muscle  by  splanch- 
nic stimulation,  101 ;  neuro- 
muscular, lessened  by  fa- 
tigue, 114-117,  120;  when 
lowered,  restored  slowly  by 
rest,  119;  when  lowered, 
restored  quickly  by  adre- 
nin,  119-123,   195. 

"Jumpers" :  exhibition  of  en- 
durance by,  223. 

Mania :  endurance  in,  222. 

Martial  virtues:  claims  for, 
by  militarists,  287  ;  import- 
ance of  preserving,  290- 
291 ;  preserved  in  competi- 
tive sports,  297-299. 

Metabolites:  influence  of,  on 
muscular  contraction,  104; 
action  of,  opposed  by  adre- 
nin, 129;  increase  adrenal 
secretion,  206-208. 

Militarists:  emphasis  of,  on 
strength  of  fighting  in- 
stincts, 286-288;  claims  of, 
as  to  values  of  war,  287; 
support  for  claims  of,  287. 


Muscle:  weakness  of,  after 
removal  of  adrenal  glands, 
81 ;  improved  contraction 
of,  after  injection  of  adre- 
nal extract,  82 ;  fatigue  of, 
84 ;  method  of  recording 
fatigue  of,  85-86;  fatigue 
of,  lessened  by  splanchnic 
stimulation,  89-93;  con- 
traction of,  when  fatigued, 
improved  by  increased  arte- 
rial pressure,  97-102 ;  irri- 
tability of,  when  fatigued, 
increased  by  splanchnic 
stimulation,  101 ;  contrac- 
tion of,  when  fatigued,  less- 
ened by  decreased  arte- 
rial pressure,  102-104;  ex- 
planation of  effects  of  va- 
ried arterial  pressure  on 
fatigued,  104-106 ;  irritabil- 
ity of,  decreased  in  fatigue, 
114-117,  120;  decreased  ir- 
ritability of,  slowly  re- 
stored by  rest,  117-118,  and 
quickly  restored  by  adre- 
nin, 119-123 ;  contraction 
of  fatigued  denervated,  in- 
creased by  adrenin,  130; 
point  of  action  of  adrenin 
in,  128-133;  use  of ,  in  strug- 
gle, 189;  energy  of,  from 
carbonaceous  material,  190- 
193;  disappearance  of  gly- 
cogen from,  190;  increased 
efficiency  of,  with  increase 
of  blood  sugar,  1*92-193; 
utility  of  adrenin  in  less- 
ening fatigue  of,  194- 
195 ;  efficiency  of,  increased 
by  distribution  of  blood  in 
pain  and  excitement,  201. 

Music :  stimulating  influence 
of,  227;  influence  of  mar- 
tial, 228. 

Neurones,  autonomic:  exten- 
sive  distribution   of    sym- 


310 


INDEX 


pathetic,  26;  arrangement 
of  sympathetic  for  diffuse 
action,  28;  restricted  dis- 
tribution of  cranial  and  sa- 
cral, 29;  arrangement  for 
specific  action,  30. 

Olympic  games:  as  physical 
substitutes  for  warfare, 
297-298. 

Operations:  in  light  anes- 
thesia hasten  coagulation 
of  blood,  174-177. 

"Ordeal  of  rice,"  9. 

Pain :  disturbing  effect  of,  on 
digestion,  18-19;  as  occa- 
sion for  adrenal  secretion, 
59-62,  63;  glycosuria  in, 
69-70;  influence  of,  on 
distribution  of  blood  in 
body,  108;  hastens  coagu- 
lation of  blood,  172-177; 
reflex  nature  of  responses 
in,  185-187;  associated 
with  action,  189 ;  stimulat- 
ing and  depressive  effects 
of,  189. 

Pancreatic  juice :  flow  of,  in- 
hibited by  excitement,  13. 

Philippine  Islands :  substi- 
tution of  sports  for  war- 
fare in,  297. 

Power :  the  feeling  of,  229. 

Psychic  secretion:  of  gastric 
juice,  5-8,  11;  of  saliva,  6; 
dependent  on  cranial  auto- 
nomic innervation,   31. 

Psychic  "tone" :  of  gastro-in- 
testinal  muscles,  13. 

Racing:  relation  of  excite- 
ment and  power  in,  221. 

Rage :  relation  of,  to  fear, 
275 ;  transformation  of 
other  emotions  into,  276; 
bodily  changes  in,  like 
those  in  fear,  276-277 ;  im- 
portance of,  as  a  fighting 
emotion,  286. 


Reflexes :  "purposive"  char- 
acter of,  185-186. 

"Reservoirs   of   power,"   216. 

Respiration:  Utility  of  in- 
creased, in  pain  and  ex- 
citement, 202 ;  value  of 
forced,  in  lessening  dis- 
tress,  203. 

Rest:  restores  irritability 
lessened  by  fatigue,  117- 
118. 

Sacral  autonomic  division : 
functions  of,  in  mechan- 
isms for  emptying,  32-34; 
activities  of,  suppressed  by 
activities  of  sympathetic 
division,  270-272. 

Saliva :  psychic  secretion  of, 
6 ;  importance  of,  for  taste, 
6 ;  flow  of,  inhibited  by  ex- 
citement, 9. 

Salivary  glands:  innervation 
of,  by  cranial  autonomic, 
31. 

^'Second  wind":  explanation 
of,  210. 

Sex:  instinct  of,  suppressed 
by  fear  and  anger,  271. 

*'Sham  feeding,"   5. 

Splanchnic  nerves:  stimula- 
tion of,  causes  adrenal  se- 
cretion, 41-43;  method  of 
stimulating,  87-88;  stimu- 
lation of,  improves  contrac- 
tion of  fatigued  muscle, 
89 ;  stimulation  of,  hast- 
ens coagulation  of  blood, 
162-167,  but  not  in  absence 
of  adrenal  glands,  167-171 ; 
severance  of,  stops  rapid 
coagulation  following  ex- 
citement, 180-182 ;  eating 
after  severance  of,  240. 

Sports :  relation  of  excite- 
ment and  power  in,  219- 
221,  296;  as  physical  sub- 
stitutes for  warfare,   297- 


INDEX 


311 


301;  moral  values  of,  300. 

Stomach:  psychic  tonus  of, 
13;  contractions  of,  inhib- 
ited by  excitement,  14-15, 
17,  and  by  pain,  19;  in- 
nervated by  sympathetic 
neurones,  27,  and  by  cra- 
nial autonomic,  31 ;  refer- 
ence of  hunger  sensation 
to,  245-247 ;  emptiness  of, 
not  the  cause  of  hunger, 
248;  contractions  of,  when 
empty,  251-253;  method  of 
recording  contractions  of, 
255-256 ;  contractions  of, 
when  empty,  associated 
with  hunger  sensations, 
256-259;  function  of  con- 
tractions of  empty,  263- 
264. 

Strength :  feats  of,  related  to 
great  emotion,  217-218, 
229. 

Sugar:  in  blood,  66,  73;  in 
urine,  69-76;  relation  of 
adrenal  glands  to,  in  blood, 
77;  increase  of,  in  blood, 
does  not  hasten  clotting, 
159,  170;  utility  of,  when 
increased  in  blood,  188- 
193;  a  source  of  muscular 
energy,  191-193;  a  means 
of  increasing  muscular  effi- 
ciency, 192-193;  use  of,  in 
body,  not  checked  by  adre- 
nin,   197-199. 

Swallowing:  inhibits  hunger 
sensation,  254. 

Sweating:  value  of,  in  emo- 
tion and  pain,  203. 

"Sympathetic"  autonomic  di- 
vision: extensive  distribu- 
tion of  neurones  of,  26; 
arranged  for  diffuse  ac- 
tion, 28;  antagonistic  to 
cranial  and  sacral  di- 
visions, 34-36;  active  in 
pain  and  strong  emotion. 


36;  emotions  expressed  in, 
opposed  to  those  expressed 
in  cranial  and  sacral  di- 
visions, 268-272 ;  domi- 
nance of,  temporary,  273. 

Threshold  stimulus :  as  meas- 
ure of  irritability.  111; 
method  of  determining, 
111-114;  increased  in  fa- 
tigue, 114-117,  120;  when 
increased,  slowly  restored 
by  rest,  117-118,  and  quick- 
ly restored  by  adrenin, 
119-123. 

Trial  by  battle:  feats  of  en- 
durance in,  226. 

Vagus  nerves:  severance  of, 
does  not  abolish  appetite, 
240-241,  and  does  not  abol- 
ish hunger  contractions  of 
the  stomach,  261. 

Viscera :  similar  changes  in, 
in  various  strong  emotions, 
275-279;  changes  in,  not 
distinctive  for  emotions, 
280-281. 

Vomiting:  in  consequence  of 
pain,  19. 

Warfare :  as  an  expression  of 
strong  emotions,  286 ;  phys- 
ical and  moral  values 
claimed  for,  287;  barbari- 
ties of,  and  opposition  to, 
289-290;  moral  substitutes 
for,  292-293;  physical  sub- 
stitutes for,  293-297;  con- 
trast between  ancient  and 
modern,  294-295. 

Witnesses :  stimulating  in- 
fluence of,   227. 

Work :  effect  of,  on  neuro- 
muscular irritability,  117; 
done  with  use  of  car- 
bonaceous material,  190- 
193. 

(1) 


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